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.AJfl 
I 



OCCASIONAL PAPERS 

OF THE 
CALIFORNIA ACADEMY OF SCIENCES 



No. 150 



May 4, 2001 



EMBIA 

Contributions to the Biosystematics 

of the 
Insect Order Embiidina 

Part 3 
The Embiidae of the Americas 

(Order Embiidina) 

By 

Edward S. Ross 




Published by the California Academy of Sciences 
San Francisco, California 



EMBIA 

Contributions to the Biosystematics of the 
Insect Order Embiidina 






#'-^- 



n 



Brachyplerembia moreliensis Ross (Embiidae), male paratypeoft' ••'' ''^^..■ 
microscope slide. Shows short wings, one of the characteristic's "** 
of the species. Some paratypes have, even shorter wings, 
scarcely the length of the bearing somite. 
Type locahty: near Morelia, Michoacan, Mexico. 



OCCASIONAL PAPERS 

OF THE 
CALIFORNIA ACADEMY OF SCIENCES 



No. 150 



May 4, 200] 



EMBIA 

Contributions to the Biosystematics 

of the 
Insect Order Embiidina 

Part 3 
The Embiidae of the Americas 

(Order Embiidina) 




By 

Edward S. Ross 



Published by the California Academy of Sciences 
San Francisco, California 



SCIENTIFIC PUBLICATIONS 

Alan E. Leviton, Editor 

Katie Martin, Managing Editor 

© 2001 California Academy of Sciences, Golden Gate Park, San Francisco, California 941 18 

All rights reserved. No part of this publication may be reproduced or transmitted in any form or by any 
means, electronic or mechanical, including photocopying, recording, or any information storage or retrieval 
system, without permission in writing from the publisher. 

ISSN 0068-5461 



Table of Contents 

Part 3. The Embiidae of the Americas (Order Embiidina) 

Abstract 1 

Introduction 1 

Checklist of American Embiidae 2 

Key to Subfamilies of American Embiidae 3 

Archembiinae Ross 3 

Key to Genera of Archembiinae 4 

Genus Archembia Ross 4 

Key to Species of Archembia 5 

Genus Calamoclostes Enderlein 17 

Key to Species of Calamoclostes 18 

Microembiinae Ross 23 

Scelembiinae Ross 24 

Key to New World Genera of Scelembiinae 25 

Genus Lithembia Ross 26 

Genus Embolyntha Davis 26 

Genus Xiphosembia Ross 28 

Genus Dolonembia Ross 30 

Genus Ischnosembia Ross 32 

Genus Gibocerciis Szumik 34 

Key to Species of Gibocerciis, subgenus Gibocercus 35 

Genus Gibocercus (Amazonembia) Ross 39 

Key to Species of Gibocercus, subgenus Amazonembia 39 

Genus Pararhagadochir Davis 43 

Key to Species of Pararhagadochir 44 

Genus Litosembia Ross 58 

Genus Biguembia Szumik 60 

Genus Argocercembia Ross 63 

Genus Aphanembia Ross 64 

Genus Ambonembia Ross 66 

Genus Malacosembia Ross 70 

Genus Ochrembia Ross 73 

Mesoamerican and Mexican Scelembiinae 75 

Genus Neorhagadochir Ross 75 

Subgenus Drepanembia Ross 75 

Genus Brachypterembia Ross 76 

Genus Conicercembia Ross 76 

Incertae Sedis 79 

Embia (Olyntha) mulleri Hagen 79 

Pachylembiinae Ross 81 

Genus Pachylembia Ross 81 

Key to Species of Pachylembia 81 

Literature Cited 84 




Parts 

The Embiidae of the Americas (Order Embiidina) 



Abstract 

All 65 recognized American species of the fam- 
ily Embiidae (30 new in this paper) are treated, as- 
signed to twenty-one genera (9 new in this paper) 
and placed in four new subfamilies. In many cases 
the treatment of species is limited to those serving 
as the type species of a genus. Additional species in 
my collection (CAS), many undescribed, will be 
treated in future works. Included will be several col- 
lected in northeastern Brazil, males of which did not 
mature in cultures in time for incorporation in this 
treatment. The arrangement of genera in this check- 
list do not always reflect degrees of relationship. 

Introduction 

Embiidae, one of the order s major families, is 
well represented in the Americas, but most species 
belong to one subfamily, Scelembiinae, which is 
more richly represented in equatorial Africa, often 
by genera closely related to those in the Americas. 
Several other embiid subfamilies are present in 
Africa and India but the family appears to stop in 
eastern Burma. Most southeastern Asian and Aus- 
tralian families are very distinct. 

My background for this paper includes a study 
of the primary types of almost all New World 
species of Embiidae, as well as those of other taxa. 
I have also visited almost all Latin American coun- 
tries and have cultured field-encountered speci- 
mens. 

Except for obvious close relationships within 
the newly-named subfamily Archembiinae, and its 
affinity to the plesiomorphic family Clothodidae, 
the development of a cladogram for other genera 
would be premature. Many anatomical characters 
cannot be well expressed in words. Also, I have late- 
ly become impressed by the importance of col- 
oration, especially that of females, in distinguishing 
species. Very often species are almost indistinguish- 
able on the basis of terminalia characters. 

The distinct embiid genera of Mesoamerica, es- 
pecially of Mexico's Sierra Madre Occidental, ap- 
pear to have evolved during the long separation of 



Mesoamerica from South America. One Mexican 
genus, Pachylemhia Ross, is so distinct, including 
copulation methods, that I propose a new subfami- 
ly, Pachylembiinae, for it. 

Explanation of terminalia symbols: 9 = ninth 
abdominal tergite; 10 L and 10 R = hemitergites of 
tenth somite; 10 LP and 10 RP = processes of these 
hemitergites; MS = medial sclerite of 10; EP = 
epiproct (somite II); H = hypandrium (sternite 9); 
HP = process of H; LPPT and RPPT = left and right 
paraprocts; LCB and RCB = left and right cercus- 
basipodites; LC, = basal segment of left cercus. 

Institutional symbols: AMNH - American 
Museum of Natural History, New York; BMNH - 
British Museum, Natural History, London; CAS - 
California Academy of Sciences, San Francisco; 
IHNB - Instituto de Historia Naturales, Bogota; 
IFML - Instituto Miguel Lillo, Tucuman; MHNP - 
Museo Historia Naturales Lima Peru; MNRJ - 
Museu National Rio de Janeiro; MZUSP - Museu 
do Zoologia de Universidad de Sao Paulo; UNAM — 
Instituto Biologia, UNAM, Mexico, DF; NMQ - 
Museo de Ciencias Naturales, Quito; USNM - 
United States National Museum, Washington. 

Nomenclature of types: For a holotype, I 
choose the best quality adult male specimen from 
the largest series with the most precise locality and 
biological data. In the case of the few partheno- 
genetic species, the holotype is an adult female. 

In accordance with my personal opinion (Ross, 
1956) that paratypes should as near as possible rep- 
resent characters of the holotype and not necessari- 
ly the general nature of the species (e.g., variation, 
distribution, etc.), my paratypes are topotypic and of 
the same sex as the holotype. My designations allo- 
type and parallotype are similarly restricted but, be- 
cause they are not paratypes, they have no formal 
status under the International Code of Zoological 
Nomenclature (4th edition. 2000). 

In most cases, my type series are from a single 
culture. Specimens apparently conspecific but from 
other localities are treated as "other specimens ex- 
amined." 



OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES 



CHECKLIST OF AMERICAN EMBIIDAE 



Subfamily Archembiinae, new 

Genus Archembia Ross 
batesi (McLachlan). Amazon Basin 
peruviana n. sp., Peruvian montana 
lacombea Ross, eastern Brazil 
kotzbaiieri (Navas), eastern Brazil 
bahia n. sp., Bahia, Brazil 
Jilata n. sp., southern Brazil 
paranae n. sp.. southern Brazil 
boliviana n. sp., SE Bolivia 
avida n. sp.. S-coastal Ecuador 

Genus Calamoclostes Enderlein 
albistriolatus Enderlein, Bafios, Ecuador 
gurneyi Ross, southern Colombia 
auriceps n. sp., central Ecuador 
micropterus n. sp., central Ecuador 
silvestris n. sp.. El Oro. Ecuador 
oculeits n. sp., W of Call, Colombia 
Subfamily Microembiinae, new 

Genus Microembia Ross 
nigosifrons Ross, Iquitos, Peru 
Subfamily Scelembiinae, new 

Genus Lithembia Ross 
florissantensis (Cockerell), Oligocene 

Genus Embolyntha Davis 
brasiliensis (Gray). Rio region, Brazil 

Genus Xiphosembia, new 
ainapae n. sp., Amazonia, Brazil 

Genus Dolonembia. new 
tapirape n. sp., Mato Grosso, Brazil 

Genus Ischnosembia, new 
amazonica n. sp.. Lower Amazon, Brazil 
surinamensis (Ross), Surinam 

Genus Gibocercus Szumik 
chaco Szumik. Argentina 
urucumi Szumik. Mato Grosso, Brazil 
beni Szumik, Bolivian yungas 
penivianus n. sp., Peru\'ian montaiia 
magnus n. sp., Bolivian yungas 

Subgenus Amazonembia. new 
sandrae n. sp.. Napo region. Ecuador 
napoa n. sp.. Napo region, Ecuador 
nanai Szumik, Iquitos region, Peru 
flavipes n. sp.. Iquitos region. Peru 

Genus Pararhagadochir Davis 
thniratis (Saussure) Trinidad, Venezuela 

= flavicollis Krauss. Rio Orinoco 
flavicollis (Enderlein), Bolivia. Paraguay 
= schadei Ross, n. syn. 



pallida n. sp., Argentina 
picclma n. sp., Machu Picchu, Peru 
trachelia (Navas), Argentina 
bicingillata (Enderlein), Amazonia 

= davisi Ross, n. syn. 
christae Ross, lower Amazon, Brazil 
balteata Ross, Sao Paulo, Brazil 
tenuis (Enderlein), Bolivian yungas 
birabeni (Navas), Argentina 
confusa Ross, Argentina 
minuta n. sp., northeastern Brazil 

Genus Litosembia, new 
oligembioides n. sp., Belem, Brazil 

Genus Biguembia Sztmiik 
multivenosa n. sp.. northeastern Brazil 
copo Szumik, Argentina 
cocum Szumik, Mato Grosso. Brazil 

Genus Argocercembia, new 
giiyana n. sp., Guyana 

Genus Aphanembia new 
ohscura n. sp.. Amazon Basin 

Genus Ambonembia, new 
incae, n. sp., Cuzco, Peru 
adspersa (Enderlein), Bolivia 

Genus Malacosembia, new 
tucumana n. sp., Argentina 
yungae n. sp., Bolivia 

Genus Ochrembia, new 
wagneri (Navas), Argentina 

Genus Neorhagadochir Ross 
inflata Ross, Guatemala 
Subgenus Drepanembia Ross 
salvini (McLachlan), Guatemala 

Genus Brachypterembia Ross 
moreliensis Ross, Michoacan, Mexico 

Genus Conicercembia Ross 
tepicensis Ross, Tepic, Mexico 
septentrionalis (Marino-Marquez), 
Michoacan, Jalisco, Colima, Mexico 

Insertae sedis 
Embia mulleri Hagen, southeastern Brazil 
Subfamily Pachylembiinae, new 

Genus Pachylembia Ross 
chapalae, Ross, Michoacan, Mexico 
audanae n. sp., Jalisco, Mexico 
colimae n. sp., Colima. Mexico 
unicincta Ross, Jalisco, Mexico 
taxcoensis Ross, Taxco, Gro, Mexico 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



KEY TO SUBFAMILIES OF 
AMERICAN EMBIIDAE 

(Males) 

1. Basal segment of left cercus (LC,) without an 
inner lobe. Left tergal process (10 LP) minute, 
needle-like. Sierra Madre Occidental, Mexico 
Pachylembiinae 

— LCi with inner, usually echinulate, lobe. 10 LP 
conspicuous 2 

2. Anterior margin of clypeus rugose, its outer an- 
gles projected. Small, upper Amazonian 
species Microembiinae 

— Margin of clypeus smooth, not projected 3 

3. Medial flap (MF) usually elevated, if not, a 
large sclerite is present in caudal membrane of 
tergite 9 Archembiinae 

— Medial flap never elevated, no sclerite in caudal 
membrane of tergite 9 Scelembiinae 

Archembiinae Ross 

new subfamily 

Type genus. — Archembia Ross, by present des- 
ignation. 

Distribution. — South America. 

Diagnosis. — Males: Size medium to large (body 
length 11-18 mm), usually winged but apterous or 
subapterous in at least two high Andean species. 
Mandibles usually short, thin, apical teeth on a sin- 
gle plane; mentum prominent; submentum quad- 
rate, weakly sclerotized, anterior and lateral mar- 
gins not inflexed. Wings at times basally broad; ve- 
nation embioid but with MA unforked as a variation 
in Archembia arida n. sp., and Calamoclostes 
oculeus n. sp.; veins well sclerotized except for 
Cuja which is weak, never forked. Hind basitarsus 
with a usually small, medial papilla. Terminalia in 
Archembia small; cerci, especially the apical seg- 
ment, often greatly elongated; terminalia of Calam- 
oclostes more robust; cerci shorter. Tenth tergite 
broadly cleft to base; medial flap (MF) elevated, 
ridge-like in Archembia; epiproct sclerite (EP) 
broadly attached to MF, narrowed caudad and slant- 
ed downward instead of horizontal. Left hemitergite 
(10 L) weakly margined on inner side; its process 
(10 LP) slanted mesad; narrow, simple in Archem- 
bia, but slightly barbed distally and dorsally in 
Calamoclostes; right process (10 RP) at most a 
short, triangular lobe, membranous on inner side, 
not terminated as a sclerotic talon. Hypandrium (H) 
uniformly but weakly sclerotized; its lobe (HP) 



short, its caudal and left margin membranous. Left 
paraproct (LPPT) composed of a membranous, set- 
ose lobe and a sclerotic outer portion toward base of 
left cercus (in /). arida, LPPT is completely sclero- 
tized, the setose portion obscure). Right paraproct 
(RPPT) reduced to an inconspicuous, setose lobe 
partially obscured beneath apex of HP. Right cer- 
cus-basipodite (RCB) a peculiar, sclerotic ring 
around base of right cercus, never fused to adjacent 
structures. Basal segment of left cercus (LC,) al- 
ways bearing a prominent medial or distal echmu- 
late lobe. 

Females. — Subfamily characters not investigat- 
ed. 

Discussion. — This exclusively South American 
subfamily contributes much to an understandmg of 
the evolution of the abdominal terminalia of males 
of the order. The medial flap (MF) is particularly in- 
teresting because it is undoubtedly homologous to 
the upturned apex of the tenth tergite of Clothoda 
Enderlein, the most plesiomorphic genus of the or- 
der. In this structure, which includes a vestige of 
somite eleven, archembiines are more closely relat- 
ed to Clothoda than that genus is to its congeners 
(Cryptoclothoda Ross, Antipaluria Enderlein, and 
especially Chromatoclothoda Ross) in which MF 
isn't upturned, having leveled off to become the in- 
ner portion of the right hemitergite (10 R) and its 
process ( 10 RP). 

Archembiines also indicate that the epiproct 
sclerite (EP) throughout the order is derived from 
the ventral (or mesocaudal) surface of MF (= ves- 
tige of tergite 1 1 ). This sclerite hinges and usually 
levels off on a supra-anal pad (EP). This pad and its 
sclerite (when present) apparently is attachment for 
intertergal muscles which pull the right process (10 
RP) downward during copulation, especially in 
oligotomoid and teratembioid genera. It should be 
noted that 10 RP has no muscles. Therefore, the 
epiproct is of great copulatory importance through- 
out the order. However, as in some genera of 
Scelembiinae, the epiproct, as well as the medial 
flap, is scarcely discernable. 

It is noteworthy that Clothoda and Archembia 
have in common the largest number of chromo- 
somes as yet observed in the order (Ross, unpub- 
lished data). 

The subfamily's two genera are distinguished by 
overall dissimilarity of male characters some of 



OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES 



which aren't exclusive, e.g.. generally more slender 
body form of Archembia with longer cerci and an- 
tennae; smaller, narrower head of Archembia with 
thinner mandibles and finer apical dentation; a usu- 
ally broader vannal hind wing area of Archembia; 
universal absence in Archembia of a large sclerite in 
the dorso-caudal membrane of the ninth abdominal 
somite; universal simplicity of left tergal process 
(10 LP); and the usually medial position of the left 
cercus lobe of Archembia. The left cercus lobe often 
is very large in Calamoclostes. The geographic 
ranges of the two genera are almost allopatric and 
colony formation and habitats usually are generical- 
ly distmct. Colonies of Archembia usually are con- 
spicuous on road bank, and bark surfaces, those of 
Calamoclostes are obscurely concealed in rock and 
bark crevices. 

Distribution. — Archembia is a prominent genus 
in Amazon and Parana river basins, and especially 
in Atlantic forest zones of eastern Brazil. Calamo- 
clostes is almost entirely confined to high Andean 
habitats in Ecuador and Colombia. However, 
Archembia arida n. sp., is isolated along the arid 
southern coast of Ecuador. It is very unlikely that it 
was introduced into the region in human commerce. 

KEY TO GENERA OF ARCHEMBIINAE 

(Males) 

1 . Aduh males with a large sclerite in posterior in- 
tersegmental, dorsal membrane of abdominal 
somite nine. Left tergal process (10 LP) with an 
acute apical hook and, at times, a blunt subapicai 
dorsal spur. Andean altiplano and coastal Ecuadori- 
an distribution Calamoclostes 

— Adult males lacking an intersegmental sclerite 
on somite nine. Left tergal process (10 LP) simple, 
apex rounded. .-Absent in Andean altiplano. Wide- 
spread in Amazon and Parana basins as well as in 
eastern Brazil. One species in arid coastal Ecuador 
Archembia 

Genus Archembia Ross 

Archembia Ross. 1971:30. Szumik. 1996:51; 1998:141 
(in cladogram). 

Type species. — Archembia lacombea Ross, by 
original designation. 

Distribution. — South .America: Entire Amazon 
and Parana basins, eastern Brazil, southward at least 
into southern Santa Catarina, and westward to the 



lower east Andean slopes of Bolivia, Peru, Ecuador 
and Colombia. One exceptional species occurs in 
arid, coastal Ecuador. 

Diagnosis. — Males: Size medium to large 
(body length 12-18 mm) body usually slender, 
somewhat dorso- ventral ly flattened, always winged. 
Head usually disproportionately small and narrow. 
Apical antennal segments usually brown but 
abruptly pure white in two species (always in A. 
batesi. usually in A. bahia). Mandibles small, flat, 
outer margins evenly arcuate; apical dentation often 
minute, finely acute; middle tooth of left mandible 
at times smaller than others. Submentum never 
heavily sclerotized but with sides well defined, an- 
terior margin never inflexed. Wings occasionally 
broad in basal, subcostal and vannal areas; RBS 
rather narrow, other veins reduced toward apices; 
Cu|a reduced to a row of setae; MA usually 
branched; cross-veins few in number; hyaline 
stripes very narrow their margins irregular Hind 
basitarsus with a medial, often small, ventral papil- 
la. Terminalia small, weakly sclerotized; ninth and 
tenth tergites short, transverse; ninth tergite mem- 
branous except along basal margin and sides, lack- 
ing a sclerotized area in caudal intersomital mem- 
brane. Tenth tergite largely membranous medially, 
almost to basal margin. Left hemitergite (10 L) 
small, inner margin weakly sclerotized, its process 
(10 LP) simple, narrow, slanted meso-caudad, at 
times flared or twisted distad. Right hemitergite (10 
R) usually produced caudad as a short acute point 
(10 RP) which is membranous on inner side. Medi- 
al flap (MF) well developed, elevated, slightly re- 
curved and microspiculate on anterior end; ventral 
surface sclerotic, wrinkled tapered caudad, extend- 
ed on surface of epiproct (EP). Ninth sternite, or 
hypandrium (H) broadly lobed (HP). Left and right 
paraprocts (LPPT and RPPT) each represented by a 
fleshy, inner, setose, membranous lobe (the left of 
which IS larger) on either side of HP and by a dark, 
sclerotic fragment, lacking a process, at base of left 
paraproct. Right cercus-basipodite (RCB) a narrow, 
dark, sclerotic ring around cercus base. Cerci very 
elongate, slender, distal segments longer than the 
basals. Inner side of basal segment of left cercus 
with an abrupt, medial, or distal echinulate lobe. 
Both cerci completely, but not always heavily scle- 
rotized (i.e., outer surfaces not membranous). 

Females: Rather large; strongly, dorso-ventrally 
flattened; legs stout. Cranium circular; pale to dark- 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



ly pigmented; if dark, a large, transverse, dorsal, 
pale spot usually present between eyes. Antermae 
22 to 24-segmented, apical segment dark; five sub- 
apical segments contrastingly white; all other seg- 
ments dark brown. Pronotum rather large, at times 
paler than other thoracic scuta, or as dark. All other 
thoracic and abdominal scuta darkly pigmented and 
evenly sclerotized; thoracic and abdominal pleu- 
rites and stemites weakly sclerotized, usually pale. 
Legs entirely pale or partially pigmented; hind ba- 
sitarsus short with a small to large, medial, ventral 
papilla. Paragenital stemites without prominent 
structures, ninth sternite emarginated medially; 
pigmentation variable. Cerci normal, usually even- 
ly pigmented, but distals may be paler in some 
species. 

Discussion. — The submentum of males is par- 
ticularly interesting. Its broad shape in combination 
with a short, weak ventral bridge appears to be ple- 
siomorphic. Some species of the genus have many 
of the long setae of the submentum clumped in par- 
allel, lateral lines. A white secretion coagulates in 
the midst of these when a specimen is killed in al- 
cohol. Close examination reveals indistinct, circu- 
lar pores between the setal sockets. These must be 
the source of the secretion. This fluid perhaps is as- 
sociated with mating for 1 have observed males 
rubbing the venter of their head on the dorsum of 
the female's head just prior to copulation. Conceiv- 
ably this activity might make the much stronger fe- 
male receptive to copulation. It might be physio- 
logically significant that the pale cranial spot is di- 
rectly over the brain. 

In some populations of Archembia the wings 
are exceptionally broad at the subcosta's base and 
in the vannal area. The latter tends to be broadest in 
the hind wing and must be a vestige of an extensive 
vannus found in many orthopteroid insects and 
their fossilized ancestors. 

Component species. — Archembia potentially is 
a large genus with species difficult to recognize on 
the basis of male terminalia characters which are so 
useful in other genera. Also, unlike most other gen- 
era, more than one species of the genus may occur 
in the same locality and habitat. Fortunately, col- 
oration characters, especially of adult females, are 
very useful and consistent. For example, the anten- 
nae of male A. hatesi (McLachlan), the only wide- 
spread species in the Amazon Basin, invariably has 
white distal segments. All other Archembia have 



distally brown antennal apices except for a minori- 
ty of specimens oiA. bahia n. sp. The pigmentation 
of the head, acrotergites, and cerci are among the 
many characters useful for identification of adult 
females. 

I have collected additional new species in 
northeastern Brazil, one in forests on mountains in- 
land from Recife. Males of one species is jet black 
with a brilliant orange head. Females are entirely 
black. At least two other new species occur under 
stones and in adjacent leaf litter in arid thornbush- 
cactus habitats of northeastern Brazil, as well as in 
similar zones of coastal Ecuador. Therefore, 
species of the genus have a wide ecological range 
from semi-desert to wet rainforest. 

All species are highly gregarious, often with 
broods of several females apparently sharing a 
large labyrinth of galleries. Colonies may exten- 
sively cover bark and road bank surfaces. The silk 
is conspicuously white, not being covered with pul- 
verized debris. 

KEY TO SPECIES OF ARCHEMBIA 

(Adult males and females) 

1. Occurring in semi-arid, coastal habitats of 
southwestern Ecuador. Left paraproct (LPPT) 
of males almost entirely sclerotized. Subapical 
inner margin of left mandible with an arcuate 
flange arida 

— Occurring east of the Andes. Left paraproct al- 
most entirely membranous, sclerotic portion 
small confined to extreme left (next to left cer- 
cus base). Inner margin of left mandible with- 
out a flange 2 

2. Occurring in Amazon Basin 3 

— Occurring elsewhere in South America 4 

3. Males with distal antennal segments always 
white. Prothorax usually tan. Widespread in 
Amazon Basin at lower altitudes batesi 

— Males with all antennal segments and protho- 
rax dark brown. Occurring in central upper 
Amazon tributary zone ("montaiia") of Peru- 
vian Andes peruviana 

4. Cranium of males uniformly pale yellow. Yun- 
gas zone, Bolivia boliviana 

— Cranium of males dark brown, or patterned 
(except in one new species from NE Brazil 
which has an orange head). Eastern Brazilian 
localities 5 

5. Females with all dorsal body sclerites, includ- 



OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES 



ing a very dark chestnut brown pronotum, very 
glossy dark brown with a faint bluish luster 
Without pale intersomital thoracic bands. All 
ventral sclerites and membranous areas pale 
tan. Cranium glossy blackish brown with two 
sharply defined interocular golden spots. Sub- 
mentum pale yellow. Brazil: Rio to Sao Paulo 
lacombea 

— Females with head and body variably pigment- 
ed dorsally and ventrally, not glossy or with 
ventral sclerites contrastingly pale. Intersomi- 
tal thoracic bands pale 6 

6. Males with left tergal process (10 LP) dilated 
apically. Right process (10 RP) short, blunt, 
concealed beneath caudal margin of 10 R. 
Eastern Brazil, NE Argentina, southcentral 
Venezuela 7 

— 1 LP tapered to a usually-sharp point. 1 RP 
acute, visible from above. Eastern Brazil 8 

7. Males with distal antennal segments usually 
white. Submentum as broad as long. Cranium 
narrow, sides caudally convergent. Brazil: 
Minas Gerais and Bahia bahia 

— Distal antennal segments always brown. Sub- 
mentum broader than long. Cranium circular, 
broadly rounded, sides not caudally conver- 
gent. Type locality: Iguagu Falls region. Relat- 
ed new species or subspecies in other regions 
dilata 

8. Males and females very darkly melanized. 
Head of females jet black with a small interoc- 
ular orange "cloud." Mouthparts very dark 
brown. Medial area of pronotum piceous, ante- 
rior margin and caudal angles orange. Cervical 
membranes and thoracic pleura dark lavender 
Acrotergite 1 largely translucent white due to 
internal fat; clouded tan caudo-medially; adja- 
cent dorsal membranes cream. Mesonotum 
black, caudal margin yellowish, membranes 
lavender. Acrotergite 2 transparent tan, inter- 
somital membranes broadly cream. Forelegs 
golden brown, other legs darker. Metanotum 
and abdominal terga black, hind legs jet black. 
Brazil: Atlantic forests E of Curitiba ..paranae 

— Head of females chestnut brown, its golden 
"cloud"" large. Mouthparts pale amber. Prono- 
tum medium brown bordered with tan. 
Acrotergite I uniformly medium brown. 
Mesonotum dark brown; margins irregularly 
tan. Pale band between meso- and metathorax 
narrower than in A. paranae. All legs tan 



clouded with brown. Abdominal terga dark 

brown, pleura cream. Brazil: Rio region 

kotzbaueri 

Archembia batesi (McLachlan) 

Embia batesi McLachlan, 1877:380. 

Embia (Olyntha) batesi McLachlan, 1885:195. 

Olyntha batesi (McLachlan), Krauss, 191 1:29. 

Rhagadochiv batesi (McLachlan), Enderlein, 1912:56. 

Embohntha batesi (McLachlan), Davis, 1940b:347. — 
Ross, 1944:413.— Barth, 1954:172.— Barth and La- 
combe. 1955:67.— Lacombe, 1958a:177; 1958b:655; 
1960, p.l; 1963:393; 1965:503.— Ross, 1970:166. 

ArcJiembia batesi (McLachlan), Ross, 1971, p. 32. 

Type. — Male, on slide, McLachlan Collection, 
BMNH. Type labels: "Embia batesi, M'L.," 
"Type"(red), "Brazil Bates" (circle). It should be 
noted that although McLachlan published, "collect- 
ed by Bates on the Amazons," "Amazons," doesn't 
appear on the specimen labels. It is likely, however, 
that the type was collected at Ega (Tefe), one of 
Bates' favorite collecting localities on the upper 
Amazon. In the British Museum I have seen a dam- 
aged specimen of A. batesi labeled "Ega" (in a blue 
circle), probably McLachlan Collection, which was 
undoubtedly collected by Bates but not used as the 
type due to its damaged condition. 

Discussion. — Nothing was added to the knowl- 
edge of this comiTion species until Davis (1940b) 
redescribed its type. I have since cultured series 
from many localities in the Amazon Basin. It ap- 
pears that A. batesi is a "weed species" spread in 
commerce. Batesi isn't congeneric with ''Embia" 
brasiliensis Gray (or Griifith and Pidgeon) which, 
to date, has been collected only in Atlantic forests 
near Rio de Janeiro. Anatomical research of Barth, 
and Lacombe, cited in the above synonymy, in- 
volved specimens of Archembia lacombea, not Em- 
bolyntha batesi, as stated by them. 

Adult males of A. batesi exhibit most of the in- 
tegumental characters illustrated (Fig. 1) for the 
following new species except for narrower wing 
bases. These two species appear to be the only 
archembias in the Amazon Basin. Archembia bate- 
si is the embiid most likely to be collected in Ama- 
zonia by the non-specialist. It is the only Archem- 
bia in the region with males having white-tipped 
antennae. The distal five antennal segments are 
white, the 21st (the most distal) is apically tan. The 
prothorax usually is tan in contrast to other dark 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



brown body sclerites. The mid- and hind coxae are 
cream white. The cercus segments are exceptional- 
ly long, the lobe of the basal segment of the left 
cercus is abruptly projected and well centered. 

Records. — Brazil: Serra do Navio. Amapa. col- 
ony on small branch in rainforest (E. S. Ross) CAS 
Esprito Santo (McLachlan Collection) BMNH 
Vaupes (Igarapes), Rio Negro (D. Lacombe) CAS 
Medio Javari, Amazonas (D. Lacombe) CAS; Porto 
Planton (D. Lacombe), CAS; Rancho Grande, 62 
km S Ariquemes, Rondonia, 187 m, bark in pri- 
mary forest (E. S. Ross) CAS; Manaus, bark in for- 
est near Hotel Tropical (E. S. Ross) CAS. Colom- 
bia: Macoa, Nariiio, colonies on tree bark (E. S. 
Ross) CAS; Sta. Rosa Sucumbios, Rio San Miguel, 
int. Putomayo, 400 m, at light, 2 males (B. Malkin) 
CAS. Ecuador: 5 km N Puyo, Napo-Pastaza, 953 
m, colony on fence post (E. S. Ross) CAS; Al- 
inahui, 25 km E Puerto Napo, 475 m, (E. S. Ross) 
CAS; 15 km. N Limon, Marona-Santiago, 1010 m 
(E. S. Ross) CAS. Peru: E end Boqueron de Padre 
Abad, (E. S. Ross) CAS; Pucallpa, limbs of shade 
trees along street (E. S. Ross) CAS; Iquitos, colo- 
nies on bark of river front trees (E. S. Ross) CAS; 
Amazon Camp, Rio Momon, 97.5 m (near Iquitos) 
(E. S. Ross) CAS. 

Archembia peruviana Ross 
new species 

(Figure 1) 

Holotype. — Male, on slide, CAS. Data. — Peru: 
Cueva de la Pava, Tingo Maria, Huanuco, matured 
XI-54 (E. S. Ross and E. I. Schlinger). 

Description. — Appearance: Moderately large 
(body length 13 mm); uniformly brown throughout. 
Color details (in alcohol): Cranium chestnut brown, 
lighter at vertex, clypeal area and caudal margins 
blending darker. Eyes lavender black. Mouthparts 
concolorous with cranium; antennae brown to apex 
(without white apical segments). Thoracic sclerites 
varied shades of medium brown, except cervical 
and prosternum which are yellow tan; pleurites of 
pterothorax with faint bluish luster; all intersclero- 
tal membranes cream white (the color of internal 
tissue). All legs medium brown to tan, including 
mid and hind coxae. Wings very dark brown with 
very narrow hyaline, intervenal stripes; cross-veins 
highly variable in number and position, none be- 
hind MP; white when crossing hyaline stripes (ex- 
tensively white in many specimens). Abdomen, ex- 



cept for darker terminalia, varied shades of medium 
brown. Dimensions (on slide). — Body length 13.0 
mm.; forewing length 7.6 mm, breadth 2.1 mm. 

Important integumental characters. — As fig- 
ured almost identical to A. batesi. However, readi- 
ly distinguished by larger size and uniformly dark 
brown color, especially the dark prothorax and en- 
tirely brown antennae (lacking white distal seg- 
ments). 

Allotype. — Female, in alcohol, with holotype 
data and disposition. 

Description. — Appearance: Moderately large 
(body length 10.0 mm), generally dark chestnut 
brown with contrasting cream prothorax and legs 
and transverse spot between eyes; antennal apices 
white. Color details: Cranium dark chestnut brown 
except for a large, transverse, somewhat chevron- 
shaped cream area between eyes. Eyes black. An- 
tennal segments 1-15 dark brown, 16-20 pure 
white, 21 light brown (number complete). Mouth- 
parts uniformly brown. Cervical sclerites clear, pale 
amber. Pronotum cream with faint light brown 
clouding; prosternum medium brown; intersclerotal 
membranes cream; acrotergites and meso- and 
metathoracic sclerites dark brown, sterna lighter; 
pleural membranes dark lavender. Legs cream ex- 
cept for tan tibiae and clouding on hind femoral 
bases. Abdominal tergites dark brown, stemites 
medially tan; pleural membranes whitish; cerci 
dark brown with dark lavender joint membranes. 

Paratypes. — Numerous adult males reared from 
cultures I collected on the grounds of the experi- 
ment station at Tingo Maria, Huanuco, Peru; on 
bark of lemon trees on bank of Rio Huallaga; on 
tree trunks 4 mi. SW Las Palmas, Huanuco; and 
Cueva de la Pava, near Tingo Maria. Deposited in 
CAS, USNM, MNRJ, MHNP, and BMNH. 

Discussion. — Extensive series of adult males 
from the Tingo Maria region have the above de- 
scribed, uniformly dark brown coloration, includ- 
ing the coxae and all antennal segments. Archembia 
batesi, a very close relative, represented in my col- 
lection by hundreds of specimens from throughout 
the Amazon Basin, invariably have the prothorax of 
the male contrastingly tan and invariably white an- 
tennal apices. The eastward distribution of .4. peru- 
viana seems to stop on the east side of the last 
mountain range between Tingo Maria and the Ama- 
zonian plain. Future studies may reveal that A. 



Sc. 



OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES 



C RBS 



'\/ >l i' 'c- 




TERMINALIA 

VENTRAL 



Figure 1. Archembia peruviana Ross, n. sp., holotjpe. Type locality: Tingo Maria, Peru. Structures are very similar in 
A. balesi (McL.). Coloration is the most convenient means of distinguishing the two species. 



peruviana is a dark subspecies of A. batesi limited 
to higher altitudes of the eastern Andean foothills at 
an average altitude of 1 .000 m. 

Biological notes. — Archembia peruviana, like 
.-1. ba!esi. forms extensive, fully exposed galleries 
on the trunks, limbs and ledges in primary forest, 
especially if semi-cleared for agriculture. Colonies 
also occur on foundations of building and even on 
steel bridges, especially around bolts. 

Archembia lacombea Ross 

(Figure 2) 

Archembia lacombea Ross. 1971:33, fia. 1. 



Holotype. — Male, on slide, CAS. Data. — Bra- 
zil: Ponte Maromba, Parque Nacional do Itatiaia, 
1200 m, mamred in culture 29-V11I-1964 (E. S. 
Ross). 

Name basis. — Named after Dyrce Lacombe, 
Fimdagao Oswaldo Cruz. Rio. a personal friend 
who conducted research in the internal anatomy of 
embiids. 

Redescription. — Appearance: Moderately 
large, slender, alate; very dark brown with head 
dull yellowish. Color details (in alcohol): Cranium 
pale orange but dulled by a uniform, faint mottling 
of rust brown which increases anteriorly and be- 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 




HIND BASITARSUS 



Figure 2. Archembia lacombea Ross, holotype (re-published from Ross, 1971 ). Type locality: Parque Nac. do Itatiaia, 
Brazil. 



comes a solid dark band around eyes and antero- 
laterally between eyes and the anterior tentorial 
pits; ventral surface paler; cranial setae blackish. 
Eyes blackish lavender, narrowly pale at edges. All 
antennal segments blackish brown except for grad- 
ually tan apical segments; total 22 segments. Sub- 
mentum golden with a whitish coagulant amidst 
bases of parallel, lateral setal clumps. Sclerites of 
thorax and legs chocolate brown with darker sutur- 
al lines; membranes whitish, strongly tinged with 
lavender Wings uniformly dark brown with very 
narrow hyaline stripes; costal and radial sinus 
(RBS) margins bright salmon pink. Sclerotic por- 
tions of abdomen dark tan, membranes cream 
white, palest around the two basal stemites; termi- 
nalia, including cerci, blackish brown; membranes 
gray lavender. Dimensions: Body length 1 1.5 mm; 
forewing length 8.1 mm, breadth 2.2 mm. Impor- 
tant integumental structures as figured. 

Allotype. — Female, in alcohol, with holotype 
data and disposition. 

Redescription. — Appearance: Rather large 
(body length 13.5 mm), flattened, dark chocolate 



brown dorsally, very glossy, sheen faintly metallic; 
prothorax somewhat paler; venter and legs very 
pale; apical antennal segments white. Color details: 
Cranium almost black with a large, transverse, 
golden band between eyes, its margins diffused; 
ventral surface becoming medium brown mesad. 
Antennal segments 1-15 concolorous with crani- 
um, segment 16 tan, 17-18 pure white, 19 basally 
white, apically tan, 20 dark brown. Mouthparts 
other than mandibles cream white, including sub- 
mentum; palpi blending to light brown. Cervical 
sclerites cream white. Pronotum chestnut brown, 
glossy; prosternum cream white, tinged with 
brown; pleurites transparent amber, surrounding 
membrane whitish. Aerotergite 1 dark brown. 
Mesonotum, aerotergite 2 and metanotum glossy, 
dark chocolate brown; meso- and metathoracic 
pleura cream white, sterna and legs similar but 
lightly tinged with brown. Abdominal terga glossy 
chocolate brown; sterna 1-7 cream white; 8 dark 
brown, paler laterally; 9 basically dark amber, heav- 
ily tinged with dark brown. Abdominal pleura 
cream white. Paraprocts pale amber. Basal seg- 
ments of cerci very dark brown, the distals ma- 



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OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES 



hogany, joint membranes dark lavender brown. 
Body length 13.5 mm. 

Paratypes and parallotypes. — Reared topotypic 
adults deposited in CAS, USNM, MNRJ, MZUSP 
and BMNH. 

Additional records. — A large series of adults 
reared from stock I collected on road banks and 
bark in Parque Nac. do Tijuca (Paineiras) above 
Rio de Janeiro (CAS) agree closely with the type 
lots from Parque Nac. do Itatiaia (1000-1800 m). 
The species occurs also within Rio in the botanical 
garden and on grounds of Catholic University. 
Southwest of Rio a male was collected at Man- 
garatiba (at light) 20-1-93; also in Reserva Ruschi, 
near Santa Teresa, Espirito Santo and, unexpected- 
ly, 10 km SE Ipiaii, Bahia, 100 m. I also cultured a 
large series from the botanical garden in Sao Paulo 
(CAS). It is possible that this apparently vigorous 
species has been moved about in human commerce. 

A series from Campos, Est. Rio de Janeiro 
(CAS) collected by Dyrce Lacombe, has very dis- 
tinct females with large, entirely golden heads 
slightly clouded with brown; legs, pleura and ven- 
ter very pale; cerci as in A. lacombea, entirely dark, 
including membranes. 1 cultured a similar series 
from 20 km N of Linhares, Espirito Santo. Its males 
have apical cercus segments unusually long. Fe- 
males reared from Minas Gerais: Cacaca 
(monastery), and 20 km S Manuaco, 650 m (both 
CAS) also have head golden blending to brown to- 
ward the clypeus. 

It is possible that specimens with gold-headed 
females represent a new species. However, a de- 
tailed study of this possibility and other systematic 
questions ideally should be undertaken by a quali- 
fied Brazilian researcher with an opportunity to 
rear series from many localities. 

Dyrce Lacombe contributed extensively to 
knowledge of the anatomy and histology of la- 
combea using the name Embolyntha batesi 
(McLachlan). Most of her study specimens were 
collected near Sepetiba. 

Archembia lacombea differs from A. batesi in 
numerous coloration and anatomical characters; for 
example. A. lacombea males never have white- 
tipped antennae (A. batesi does), it has parallel 
setal clumps on the submentum (setae evenly dis- 
tributed in A. batesi). and the left cercus lobe is api- 
cal or subapical in A. lacombea (submedial in A. 



batesi); also, A. batesi appears to be restricted to 
the Amazon Basin. 

Of the closely related species or races in the Rio 
region, I am somewhat arbitrarily applying the 
name, Embia Kotzbaiieri Navas to one of these, as 
treated below. At Paineiras, in the national park 
above Rio, colonies of A. lacombea and A. 
kotzbaiieri occur on the same road banks. Strange- 
ly, few differences in the male terminalia separate 
these species but, fortunately, they are readily dis- 
tinguished by striking coloration differences, espe- 
cially in adult females. 

Archembia kotzbaiieri (Navas) 

Embia Kotzbaiieri Navas, 1925:67; Ross, 1944:498 (as 
unrecognizable). 

Archembia kotzbaiieri (Navas), Ross, 1971. P. 32 (new 
combination). 

Type. — Male, originally in Navas Collection, 
but now apparently lost. Data. — "Heim at Brazil- 
ian: Nistheroy, 25-XI-1924. In meiner Sammlung." 

The type locality undoubtedly is Niteroi, a dis- 
trict within Rio de Janeiro. Perhaps in 1924 the area 
was less congested and one or more species of 
Archembia could have existed there. Navas' type 
appears to be lost. I couldn't find it in his collection 
when it was lodged in Zaragoza (the collection has 
since been moved to Museo Zoologia, Barcelona). 
The original description is inadequate; however, 
Navas' described coloration, and head and wing il- 
lustrations, indicate that the species is an Archem- 
bia. Furthermore, all other native embiids in the 
Rio area, except for A. lacombea and very distinc- 
tive Embolyntha brasiliensis, are very small 
anisembiids and teratembiids. I decided, therefore, 
to assign the name A. kotzbaiieri to the genus 
Archembia as the second species of that genus oc- 
curring in Rio, the first being A. lacombea. The 
species is herewith redescribed from a neotype se- 
lected from extensive cultured series of both sexes. 

Neotype. — Male, on slide, MNRJ. Data. — Bra- 
zil: Parque Nac. do Tijuca (at Paineiras) above Rio 
de Janeiro, matured 25-VI-64 in culture 695A 
(E. S. Ross). 

Description. — Appearance: Moderately large 
(body length 1 1.0 mm), slender, alate, light brown 
throughout except for a cream pro-pterothoracic 
band and cream abdominal pleura. Color details: 
Cranium basically tan, becoming darker laterally. 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBllDINA, PART 3 



11 



toward clypeus and around dark lavender eyes. An- 
tennae uniformly medium brown grading to tan, but 
not white distad; 28-segmented. Submentum and 
venter of cranium very pale amber, setae of sub- 
mentum sparse, only slightly clumped in lateral 
lines, these are clogged anteriorly with a thick, 
white, coagulated secretion. Sclerites of body and 
legs varied shades of light brown and tan with dark 
brown sutural lines. Membranes white, those be- 
tween pro-and pterothorax especially conspicuous, 
forming a pale band both dorsally and ventrally; a 
whitening of the caudal margin of the pronotum 
contributes to this. Hyaline stripes of wings very 
narrow; all veins, except towards apices, sclerotized, 
except for Cu; 4 cross-veins between RBS and RP, 
between RP and MA,+2 and 1 between MA and MR 
Abdominal terga basically whitish but mottled with 
dark brown, especially toward sides and abdominal 
base; pleural membranes white; terminalia sclerites 
chestnut brown, membranes whitish; cerci dark 
brown. Dimensions (on slide): Body length (exclud- 
ing cerci) 11.0 mm; forewing length 7.5 mm, 
breadth 2.0 mm. Important anatomical structures 
similar to those of A. kicombea (Fig. 2). 

Neallotype. — Female, in alcohol, MNRJ. Data: 
A "sister" of the neotype. 

Description. — Cranium golden mahogany 
brown dorsally with very faint vertex pattern, but 
including a transverse golden "cloud" between eyes. 
Antennae brown except for white distal third of seg- 
ment 18; 19 pure white; 20 basally white, distal half 
tan; 21 entirely tan. Mouthparts, including submen- 
tum, pale amber, palpi light brown. Cervical scle- 
rites golden brown, adjacent membranes dark laven- 
der. Pronotum basally amber, tinged medially with 
dark lavender; prostemum pale amber, anterior mar- 
gins very dark lavender; pleura dark to pale amber, 
membranes whitish. Acrotergite 1 golden amber. 
Mesonotum largely medium brown, medially tinged 
with dark purple, anterior angles blending to white, 
posterior angles becoming pale amber. Acrotergite 
2, pale amber. Metanotum similar to mesonotum 
but with smaller pale angles. Meso- and metatho- 
racic pleural sclerites translucent dark amber, mem- 
branes very dark lavender; stemites translucent pale 
amber. Coxae, trochanters, and femora of all legs 
varied shade of amber; hind femora, tibiae and fore- 
tarsi tinged with lavender. Abdominal terga golden 
brown, tinged with lavender, partially bordered with 
dark lavender; sterna 1-7 clear amber medially, 



blending to dark lavender in lateral thirds; genital 
sternite 8 dark lavender, 9 basically amber but 
heavily tinged with dark brown laterally and around 
genital opening; pleurites very pale amber; mem- 
branes cream white; paraprocts grading from pale 
amber to cream caudad. Basal segments of cerci 
dark lavender brown, distal segments golden tan, 
joint membranes dark cream. Body length 16.0 
mm. 

Paraneotypes. — Numerous topotypic adults de- 
posited in CAS, USNM, MNRJ, MZUSP and 
BMNH. 

Discussion. — Although there are anatomical 
differences between males of A. lacombea and A. 
kotzhaueri, these are difficult to define except for 
the more circular head of the latter. Fortunately, the 
two species are readily distinguished by very dis- 
tinct coloration, especially that of females. Archem- 
bia lacombea is overall darker in both sexes. Fe- 
males and nymphs of ^. kotzbaiieri have a "check- 
ered" appearance, a pale cranium, acrotergites and 
body sclerites light brown, and paler cerci. In A. la- 
combea adults all these structures are much darker 
with a pronounced iridescent luster. 

Archembia bahia Ross 
new species 

Holotype. — Male, on slide, MNRJ. Data. — Bra- 
zil: On hill 20 km SW Jequie. Bahia, 600 m. colony 
in road bank just below microwave tower, matured 
in culture June 1992 (E. S. Ross). 

Description. — Appearance: Large, slender, dark 
chocolate brown except for golden head and pro- 
thorax, a whitish band between pro- and mesotho- 
rax, and usually white-tipped antennae. Color de- 
tails (in alcohol): Cranium golden with faint 
smokey pattern and black setae; eyes black; mouth- 
parts golden except for dark brown palpi (a white 
coagulant is deposited amidst laterally-clumped, 
black setae on submentum). Antennae almost as 
long as body; segments 1-19 dark brown, 20-23 
white, 24 tan. Cervical sclerites and prothorax bril- 
liant gold, setae black, membranes basically white 
but lavender-tinged on crests of folds. Membranes 
between pro- and mesothorax pure white (fonning 
a band). Coxae, trochanters and femora of forelegs 
tan, tinged with brown; tibiae and tarsi dark brown. 
All other sclerotized portions of body dark choco- 
late brown, especially the terminalia with its very 



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dark membranes; body sclerites very glossy; mid 
and hind legs lighter brown. Wing bands medium 
brown, hyaline stripes very narrow; two white 
cross- veins between RP and MAj+i, three between 
MA and MP. Dimensions (on slide): Body length 
13.75 mm; forewing length 8.0 mm, breadth 2.4 
nun. 

Allotype. — Female, in alcohol, CAS, from holo- 
type's culture. 

Description. — Appearance: Large, robust. Head, 
prothorax and acrotergite chestnut brown, otherwise 
chocolate brown except for white-tipped antennae. 
Color details: Cranium basically golden, tinged 
with chestnut brown; surface between eyes paler; 
eyes black; mouthparts light brown. Antennal seg- 
ments 1-18 brown, 19-22 white; 23 white basally, 
tan distally. Prothorax varied shades of chestnut 
brown, forelegs concolorous, cervical sclerites and 
pronotum amber; membranes heavily tinged laven- 
der. Remainder of thorax varied shades of chocolate 
brown, sternites paler, all membranes dark lavender; 
coxae, trochanter, and tarsi of mid-legs pale amber, 
femora and tibiae medium brown; hind legs rather 
uniformly brown. Abdomen glossy chocolate brown 
dorsally, paler ventrally; membranes white, tinged 
with lavender; genital sternite and cerci (including 
joint membranes) dark brown. Body length 18 mm. 

Paratypes and parallotypes. — 14 topotypic 
males and females reared in holotype s culture. De- 
posited in CAS, MZUSP and MNRJ. 

Additional record. — One male with white- 
tipped antennae from 20 km N of Linhares, Espiri- 
to Santo, 40 m. matured VII-92 (E. S. Ross) CAS. 

Discussion. — Usually males of this colorful spe- 
cies can be distinguished from all other Archembia 
in the Atlantic regions of Brazil by their white- 
tipped antennae. However, in my cultured series ten 
males have white-tipped antennae and four have 
them grading distad to light brown. Such a variation 
is unusual. 

A more consistent recognition character is the 
narrow, brilliant gold head, including all non-man- 
dibular mouthparts except brown palpi, in combina- 
tion with golden cer\ical sclerites, and largely gold- 
en prothorax, forecoxae. trochanters, and femora. 
All other sclerotized portions are dark chocolate 
brown with purple-tinged membranes. 



Archembia dilata Ross 
new species 

(Figure 3) 

Holotype. — Male, on slide, deposited in 
MZUSP. Data. — Brazil: Foz do Iguagii. Parana, ma- 
tured m culture 24-VII-1964 (E. S. Ross). 

Description. — Appearance: Large, robust, alate; 
dark brown except for bright golden head and pro- 
thorax. Color details (in alcohol): Cranium golden, 
lacking pattern, darker around antennal bases; setae 
contrastingly dark. Eyes dark lavender. Basal anten- 
nal segment dark chestnut brown, other segments to 
apex (segment 24) very dark brown. Mouthparts, in- 
cluding labrum, amber yellow except for dark 
brown palpi. Pronotum. pro-pleurites, and cervical 
sclerites yellow, slightly paler than cranium. 
Prosternum pale tan mottled with brown. Pterotho- 
racic sclerites dark chocolate brown with darker su- 
tural lines, membranes purple; legs concolorous 
with pterothorax except fore- and mid-coxae, 
trochanters, and hind trochanters which are yellow 
tan. Wings dark brown with bright pink costal and 
radial blood sinus (RBS) margins; hyaline stripes 
very narrow and weak basad; numerous cross-veins 
behind RP are white when crossing a hyaline stripe. 
Abdominal tergites brown laterally, otherwise ex- 
tensively membranous; membranes dorsally and lat- 
erally pale purple; sternites more strongly sclero- 
tized and basically dark chestnut but dark laterally 
and thus forming two longitudinal, ventral, dark 
stripes. Sclerotic portions of terminalia, including 
processes and cerci, very dark chocolate brown, cer- 
cus membranes as dark as segments. Dimensions 
(on slide); Body length 15.0 mm; forewing length 
7.6 mm, breadth 2.1 mm. 

Important integumental characters. — The large, 
robust body form. Broad head with an exceptional- 
ly large, but not heavily sclerotized submentum 
which has evenly distributed setae. Left tergal 
process narrow, parallel-sided then slightly flared 
and rounded at apex. Right process not produced, its 
apex obsolete. Hypandrium lobe scarcely produced. 
Basal segment of left cercus stout, its width through 
the lobe equal to the segment length. These and 
other features are figured. 

Allotype. — Female, in alcohol, (MZUSP) with 
holotype data. 

Description. — Appearance: Large, mostly dark 
mahogany brown with prothorax and basal half of 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORX)ER EMBIIDINA, PART 3 



13 




TERMINALIA 

VENTRAL 



Figure 3. Archembia dilata Ross, n. sp., holotype. Type locality: Foz do Igua?u, Parana, Brazil. 



fore- and mid-legs largely golden tan; apices of an- 
tennae gray white. Color details: Cranium chestnut 
brown with a narrow, suffused, golden area arcing 
caudad between antennal bases; stem of ecdysial su- 
ture also golden, as well as cranium's ventral sur- 
face; dorso-basal pattern not developed. Eyes dark 
lavender Antennae dark brown except for four api- 
cal segments which are gray white, each tipped with 
rust red; 21 segments in complete antenna. Mouth- 
parts golden except for palpi which become brown- 
ish distad. Prothoracic sclerites, including first 
acrotergite, golden with narrow brown margins and 
medial clouding on pronotum. Other thoracic ter- 
gites and pleurites dark mahogany brown; nota 
clouded with gold tan and having a distinct, pale, 
medial, longitudinal line; all stemites transparent 
pale amber, bordered with light brown; all thoracic 
membranes whitish, lightly tinged with lavender. 
Fore- and mid-legs largely golden tan basally, tibiae 
and tarsi medium brown, those of mid-legs paler; 
hind legs various shades of medium chestnut brown. 
All abdominal tergites dark mahogany brown, pleu- 



rites medium brown; sternites clear pale amber ex- 
cept sternite 8 purple white; lateral sclerites very 
dark brown; sternite 9 medium brown, paler caudad; 
paraprocts pale tan; cerci dark brown with joints 
dark lavender. Body length 17 mm. 

Paratypes and parallotypes. — Numerous reared 
topotypic adults deposited in CAS, USNM, 
MZUSP MNRJ and BMNH. 

Other specimens examined. — Several adult 
males from "Rondon 24°38'S, 54°07'W" eastern 
Parana, Brazil, XI to XII (Fritz Plaumann) CAS. I 
have collected related species or races, with a dilat- 
ed left tergal process, in other regions of Brazil and 
Bolivia but study of these should be deferred until 
ample research time and specimens are a\ailable. 
One of these, a series from Camaca, Bahia is very 
closely related to A. dilata. but has a much narrow- 
er submentum and other small distinctions. This 
also applies to a large series I reared from stock col- 
lected in Santa Elena, a Venezuela-Brazil border 
town north of Boa Vista, Roraima. Males of this 



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OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES 




SUBMENTUM 



Figure 4. Archembia arida Ross. n.sp.. holotype. Type locality: Near Santa Elena, Guayas, Ecuador. 



group of species, or races, are distinguished by the 
dilated apex of the left terga! process (10 LP); the 
small, almost obsolete right process (10 RP) ob- 
scured beneath the caudal margin of 10 R (thus not 
visible from above), the golden color of the head, 
and many minor characters. 

Biology. — The type colonies were scattered on 
the bark of trees growing along trails provided for 
tourists visiting Foz do Iguagu. The species is high- 
ly gregarious, as are all other species oi Archembia. 
The silk galleries are fully exposed lacking a cover 
of pulverized feces or debris. At the time of en- 
counter, late April 1964. most colonies consisted of 
an adult female and her brood of half-grown 
nymphs developing in unison. One colony was more 
ad\anced. howe\er. with the brood about one instar 
short of maturity. In cultures, field-collected 
nymphs matured during late July, especially during 
August, and declined during September. In the fol- 
lowins vear adults beaan maturins durins June. 



Eggs are laid on their "back" in flat clusters and 
imbedded in a matrix of pulverized material. 

Archembia paranae Ross 
new species 

Holotype. — Male, on slide, MZUSR Data. — 
Brazil: Parana Recanto Bela Vista, picnic ground 
above Joao Sao da Graciosa (between Moretes and 
PR410), 800 m. matured X-99 (E. S. Ross). 

Description. — Appearance: Slender, generally 
blackish except for the largely mottled tan cranium. 
Color details (in alcohol): Anterior two-thirds of 
cranium basically yellow tan but tinged with dark 
brown, especially on clypeus; caudal third and sides 
dark brown. Eyes jet black. Antennae entirely black, 
22 segments (complete). Venter of cranium and 
mouthparts amber except for dark brown palpi. Pro- 
thoracic sclerites and fore-legs blackish brown, 
membranes very dark lavender except narrowly 
white behind cranium. Pterothoracic scuta blackish 



ROSS: EMBIA. BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



15 



brown apically, blending to lighter brown caudad; 
acrotergite 1 and prescutum 1 blackish brown; 
acrotergite 2 mottled tan; ventral thoracic sclerites 
dark chocolate brown, sutures black. Wing bands 
dark brown; hyaline stripes narrow, margins indefi- 
nite with macrotrichia arising from dark brown 
wing spots; all cross-veins narrowly white. Abdom- 
inal sclerites dark brown except for mottled tan me- 
dial areas on somites 8 and 9; pleural membranes 
white; 10 L, 10 R and basal segments of cerci jet 
black, 10 LP dark amber, MF medium brown, ter- 
minalia membranes cream, distal cercus segments 
grading from dark brown basally to golden brown 
apically. Dimensions (on slide): body length 14.0 
mm; forewing length 10.0 mm, breadth 2.7 mm. 

Anatomical features. — Submentum as in A. bo- 
liviana very weakly sclerotized; setae very long, 
equal in length, denser in lateral areas. Terminalia 
similar to A. boliviana but 10 RP is larger and pro- 
jected meso-caudad. Lobe of left cercus narrow; 
submedial, slightly more basad. 

Allotype. — Female in alcohol, MZUSP. Data. — 
From holotypes culture. 

Description. — Appearance: Large, multicolored, 
antennal apices white. Color details: Cranium dull 
black dorsally with a conspicuous gold tan trans- 
verse band between antennal bases, gular area gold- 
en brown. Antennal seginents 1-15 dark brown, 16 
apically tan, 1 7-22 pure white, distal segment pale 
tan (23 segments, complete), mouthparts amber, 
palpi black. Pronotum golden brown bordered with 
amber, prostemum and cervical sclerites mahogany 
brown, bordered with amber; membranous areas 
dark lavender. Acrotergite 1 amber brown, adjacent 
membranes cream white. Mesoscutum glossy black, 
caudal membranes cream; pleurites black, associat- 
ed membranes dark lavender; posternum and me- 
sostemum amber, extensively dark brown. Metatho- 
rax similar to mesothorax. Fore- and mid-legs with 
coxae mahogany brown, femora amber blending 
distad to chestnut brown, tibiae dark brown, foretar- 
si dark brown, midtarsi tan; hind legs blackish ex- 
cept for tan trochanters, tibial bases and tarsi. Ab- 
dominal terga black, pleural membranes cream; 
sterna dark brown, medial area of sternite 8 and 9 
dark lavender. Paraprocts and adjacent membranes 
pale cream. Cerci, including joints, jet black. Body 
length 17.0 mm. 

Paratypes and parallotypes. — A large series 
from holotype s culture distributed CAS, MZUSP, 



MNRJ, USNM, and BMNH. Also a series from the 
same locality cultured in 1993. 

Other specimens studied. — A large culmred se- 
ries from Hacienda Belo Vista, E of Curitiba, 
Parana, 300 m, C-72 1 A + B, 1 964 (E. S. Ross). One 
pair, C-715, N of Itajai, near sea level (E. S. Ross). 

Biology. — Stock for the type cultures was col- 
lected in a colony on a mossy stump, another on the 
mossy damp surface of a concrete building, both in 
a clearing in dense cloud forest, 800 m elev. At 300 
m, east of Curitiba, colonies were on mossy stumps 
in semi-cleared pasture with scattered Awicaiia 
trees. Paratypes matured in cultures during July and, 
mostly, during August and September. Those at 300 
m mostly matured during August or October, one as 
early as March. 

Discussion. — Topotypic males are exceptionally 
large and very black. Females are multicolored with 
conspicuous pale thoracic bands. The species is 
probably widespread in cloud forests at higher ele- 
vations in Atlantic forests in Parana. One would ex- 
pect that the pair from near sea level, just north of 
Itajai would be Embia mulleri Navas, but females 
differ in having the thorax broadly cream-banded 
and the fore and midlegs are uniformly pale yellow. 
Therefore, the population is more closely related, 
perhaps racially, to A. paranae.The type of E. mul- 
leri, a female, is larger, almost entirely dull black 
with very narrow, pale thoracic band and probably 
entirely dark antennae. 

Archembia boliviana Ross 
new species 

Holotype. — Male, on slide. CAS. Data. — Boliv- 
ia: Angostura, 70 km SW of Santa Cruz matured 20- 
VIII-64 (E. S. Ross). 

Description. — Appearance: Rather large, alate. 
Cranium straw yellow. Pronotum light tan with nar- 
row brown margins, pleura and prosternum light 
brown. Remainder of body and legs uniformly ma- 
hogany brown with piceous sternal sutures; area be- 
tween pro- and mesothorax mahogany brown with 
dark membranes; pterothoracic scuta uniformly ma- 
hogany brown. Abdoinen, sclerotic portions of ter- 
minalia and cerci also uniformly mahogany browTi 
except for yellow apical half of 10 LP. Dimensions: 
Body length 12.0 mm; forewing length 8.5 rmn. 
breadth 2.1 mm. 



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OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES 



Anatomical features. — Not very distinctive. 
Cranium and mouthparts weakly sclerotized; setae 
on submentum rather evenly distributed, two espe- 
cially long setae submedially in anterior third. Cra- 
nium rather broad; eyes not inflated, separated by 
two eye-widths. Left tergal process ( 10 LP) minute- 
ly spiculate on outer surface; right tergal process 
(10 RP) minute, sharp, projected beneath caudal 
edge of 10 R. Sclerotic portion of left paraproct 
(LPPT) confined to extreme left, membranous por- 
tion extensive, clear with few setae. Lobe of left cer- 
cus large, apical. 

Allotype. — Female in alcohol, CAS. Data. — 
From holotype's culture, matured 20-VIII-64 (E. S. 
Ross). 

Description. — Appearance: Uniformly mahoga- 
ny brown with yellowish cranium, pale intersomital 
thoracic bands, and white antennal apices. Color de- 
tails: Cranium basically pale yellow with light 
brown tint, no vertex pattern. Eyes black. Antennal 
segments 1-17 mahogany brown, 18 apically white, 
19-23 white (complete). Body and legs uniformly 
mahogany brown with pronotum slightly darker and 
glossy; dorsal membranes before and caudad of 
acrotergite 1 pale, forming a faint band; comparable 
membranes between meso- and metathorax pale, 
anterior margin of metanotum broadly white, thus 
forming a conspicuous pale band, all other body 
membranes light brown only slightly paler than 
sclerotic areas. Bodv length 16. mm. 



Paratypes and parallotype. 
males, one female (CAS). 



-Two topotypic 



Discussion. — Because of the remote, far west- 
ern occurrence, and contrasting coloration of the 
head and prothorax. it is concluded that this is a dis- 
tinct species but one, as is usual within the genus 
Archembia, having only slight male terminalia dis- 
tinctions. 

Archembia arida Ross 
new species 

(Figure 4) 

Holotype. — Male on slide, CAS. Data. — Ecua- 
dor; 21 mi NE Santa Elena, Guayas. 50 m elev (est.) 
29-1-55 (E. S. Ross). 

Description. — Appearance: Moderately large, 
alate; dark brown except for bright golden head and 
white band bet^veen prothorax and pterothorax and 



whitish abdominal pleura. Color details (in alco- 
hol): Cranium almost uniformly golden, pattern 
faint. Antennae dark brown basally, becoming 
medium brown to apex, apical segments therefore 
not whitish, 19-segmented (complete). Eyes black. 
Mandibles amber yellow except for piceous inner 
margins and dentation; other mouthparts pale 
amber except for brown palpi. Body sclerites, legs, 
and cerci uniformly brown; pleural membranous 
areas whitish, terminalia membranes mottled with 
purple. Dimensions (on slide): Body length 12.0 
mm; forewing length 7.0 mm, breadth 2.0 mm. 

Important anatomical features. — Mandibles 
elongated due to lengthening of molar area, medial 
flange of left mandible much larger than in con- 
geners; submentum weak, wider than long, setae 
variable in size, evenly spaced (not clumped in lat- 
eral rows). Wings with MA forked beyond basal 
half (unforked in some specimens). Terminalia sim- 
ilar to congeners, lacking a sclerite in caudal mem- 
brane of tergite 9. As in A. dilata, cercus segments 
are stouter and shorter with the left cercus lobe ter- 
minal, broadly rounded and finely echinulated. 
Outer half of this segment desclerotized, that of the 
right cercus sclerotized only on its inner face. Left 
paraproct well sclerotized throughout; its setose, 
membranous lobe small, obscure. 

Allotype. — Female, in alcohol, with holotype 
data and disposition. 

Description. — Cranium amber yellow with rust 
brown pattern; eyes black; antennae pale tan 
throughout, 20-segmented (complete). Body scler- 
ites and legs light brown, membranous areas cream 
white. Paragenital sclerites dark brown, cerci tan. 
Body length 15 mm, somewhat distended in alco- 
hol. 

Paratypes parallotypes. — Topotypic males and 
females deposited in Museo Nacional and Catholic 
University, both in Quito, Ecuador; CAS, USNM 
and BMNH. 

Discussion. — The occurrence of an Archembia 
on the coast of Ecuador is surprising — more so be- 
cause of its occurrence in arid habitats. At first I 
thought that it might be a species of Calamoclostes, 
a genus almost entirely confined to moist, often 
high Andean habitats; however, the absence of a 
sclerite in the caudal membrane of the ninth tergite, 
the simple left tergal process and other characters, 
rules against this. The type series is from localities 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



17 



close to Santa Elena, Guayas. Here colonies were 
very conspicuous on trunks and branches of small 
trees, and amidst thorns of cacti. Most adults ma- 
tured in cultures durmg January. A large series was 
also cultured from stock collected by David Cav- 
agnaro 2 mi S of Manglaralto. Guayas with most 
adults maturing during September. He collected an- 
other culture 5 mi N of San Pablo, Guayas. Adults 
matured durmg July and August. Recently, 1 cul- 
tured the species from stock collected just south of 
Bahia, Manabi, 15 m elev., males matured during 
July. Here the habitat was seasonally-arid jungle 
with scattered silk cotton trees. It is apparent that A. 
arida has a wide occurrence in xeric habitats of 
coastal Ecuador. At higher elevations in this zone 
there are evergreen tropical forests but these haven't 
been investigated for embiids. 

In addition to their geographically separate oc- 
currence. A. arida males are distinct in their more 
elongate mandibles, almost completely sclerotized 
left paraproct, and relatively bright coloration. The 
unicolorous brown antennae of both se.xes (lacking 
white apical segments) is also distinctive. 

Genus Calamoclostes Enderlein 

Calamoclostes Enderlein, 1909:188.— Krauss, 1911: 
73.— Enderlein, 1912:27.— Navas, 1918:94.— Davis, 
1940a: 189.— Ross, 1944:414. 

Type species. — Calamoclostes albistriolatus 
Enderlein, by original designation. 

Distribution. — Ecuador and Colombia, chiefly 
in Andean altiplano. 

Diagnosis. — Males: Generally large and robust; 
alate, micropterus, or apterous; usually darkly pig- 
mented throughout, including antennae and cerci. 
Wing venation embioid, but MA isn't forked in 
some specimens of C octileus. n. sp. All characters 
similar to those of Archembia except for consistent 
differences in terminalia, as follows: Ninth somite 
with a large, sclerotized area, lacking setae, in 
medio-caudal, dorsal membrane which is slanted 
caudad. Left tergal process (10 LP) stouter, more 
sclerotic, with a distinct apical hook and often an 
obtusely-angulate, dorso-medial projection on outer 
side. Medial flap (MP) not greatly elevated; more 
transverse, microspiculate on inner-basal crest. 
Epiproct sclerite (EP) very broadly attached to MF, 
then caudally narrowed on a flatter plane than in 
Archembia (in which it is directed ventrad). Lobe of 



hypandrium (HP) membranous apically, often 
acutely tapered. Left paraproct sclerotic next to base 
of left cercus; mesal two-thirds membranous, se- 
tose, greatly enlarged, often extensively projected 
caudad; some species have a small, setose sclerite 
isolated between the mesal membranous lobe and 
sclerite at cercus base. Basal segment of left cercus 
with lobe always apical, large, the segment almost 
as broad across lobe as long; apex of lobe rounded, 
densely micro-echinulate. Right cercus-basipodite 
(RGB) not a complete ring (dorsally obsolete). 
Basal segment of right cercus with outer side often 
membranous; distal segments of cerci shorter than 
basals. 

Females: Not studied for generic level charac- 
ters. 

Discussion. — In most of its range, and entirely 
so in the high Andes. Calamoclostes species are the 
only large-sized embiids and the only representa- 
tives of the family Embiidae. Also, they are the only 
embiids in these regions having two hind basitarsal 
papillae. At lower altitudes east of the Andes, and in 
northern Colombia, it is possible that the range of 
Calamoclostes may overlap that of Antipaluria of 
Clothodidae and Pararhagadochir and Gihocerciis 
of Scelembiinae, both of which also have two hind 
basitarsal papillae. 

Species of Calamoclostes perhaps are apomor- 
phic derivatives of the more plesiomorphic genus 
Archembia . In addition to consistent anatomical dif- 
ferences between the two genera, there are distinc- 
tions in biology. Calamoclostes species, unlike 
those of Archembia, do not form extensive, sheet- 
like colonies often occupied by scores of individu- 
als. Instead, each Calamoclostes "nest" is occupied 
by a single female and her brood. Colonies are scat- 
tered, never interconnected. Some species are found 
in inconspicuous colonies obscured by bark flakes 
of trees and fence posts, others occur on road or trail 
banks at high altitude with refuge galleries extend- 
ed into soil or rock crevices. 

Component species. — 1 have collected and cul- 
tured about five species. Included are the two 
known species; C albistriolatus Enderlein from 
Baiios, Ecuador, and C gurneyi Ross ft"om southern 
Colombia. 1 also encountered an apparently new 
species far to the south in Ecuador, near Loja, but 
was unable to develop a culture. At this time 1 will 
describe only the most distinct new species. The 
genus appears to be most diversified in Ecuador's 



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OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES 



altiplano where males of some species are apterous, 
or micropterus, the one from near Paute, Azuay pos- 
sesses miniature wings (not antipenuhimate pads), 
length not exceeding the bearing thoracic somite. 
An intensive study of the genus by a qualified resi- 
dent specialist should prove to be very interesting. 

KEY TO SPECIES OF CALAMOCLOSTES 

(Males) 

1. Males apterous or short-winged. Highlands of 
Ecuador 5 

— Males with normal wings. Ecuador and Colom- 
bia 2 

2. Costal and distal wing margins narrowly white. 
Lowland rainforests of western Ecuador 
silvestvis 

— All wing margins brown. Highlands of Ecuador 
and Colombia 3 

3. Eyes very large, cranial interspace only slightly 
more than an eye-width. Highlands W of Cali, 
Colombia ocitleits 

— Eyes relatively small, cranial interspace more 
than eye-widths. Highlands of Colombia and 
Ecuador 4 

4. Right tergal process (10 RP) conspicuous, 
acutely triangulate. Membranous portion of left 
paraproct (LPPT) large, transverse. Baiios re- 
gion, Ecuador albistriolatiis 

— Right process (10 RP, inconspicuous, vestigial. 
Membranous portion of LPPT small, globose, 
projected caudad. Upper Putumayo, Colombia 
giirneyi 

5. Apterous, neotenic. Cranium largely golden. 
Ecuadorian altiplano, near Alausi aiihceps 

— Brachypterous, wings very small, not much 
longer bearing somites. Cranium uniformly 
dark brown. Ecuadorian altiplano, near Paute 
micropterus 

Calamoclostes albistriolatiis Enderlein 

(Figure 5) 

Calamoclostes albistriolatiis Enderlein, 1909:188. — 
Krauss, 191 1:73.— Enderlein, 1912:28.— Navas, 1918: 
94.— Davis. 1940a: 189.— Ross, 1944:415. 

Type. — Male, pinned: terminalia and set of 
wings on slide; deposited in Polska Academia Nauk. 
Warsaw. Labels. — "Ecuador, Baiios. 1800 m, 21-3- 
1899, E. Schmidt S." "Type" "Calamoclostes al- 
bistriolatus Endl. Type det Dr. Enderlein." 

Discussion. — To date all references to this very 



distinct species were based on Enderlein's type. 
During my first visit to Bafios (1955) I encountered 
numerous, but very obscure, colonies of this species 
in rough bark of indigenous trees (Lauraceae) along 
the town's southern street. Most collections were 
made near the Palace Hotel but such occurrence has 
probably now ceased. Recently, in the 1990s, I 
found numerous colonies on fence posts and bases 
of small trees, about 1000 m in altitude above 
Banos, on the NE slope of Volcan Tungurahua. The 
embiids occupied silk-lined retreats under loose 
bark and feed on lichens and decomposing bark. 
The silk is gray in color and inconspicuous. Each 
colony consisted of a single female with eggs or 
brood. Evasive movement is sluggish. 

This species is readily recognized by the con- 
spicuous broad white "slashes" across all cross- 
veins. These aren't as conspicuous in other species 
of the genus. This peculiarity must have suggested 
the name, albistriolatiis. Nymphs and adult females 
are characterized by variegated head, body and leg 
maculation peculiar to the species. 

Calamoclostes giirneyi Ross 

(Figure 6) 
Calamoclostes giirneyi Ko&s. 1944:415. figs. 17-19. 

Holotype. — Male, on slide, USNM. Data. — Co- 
lombia: upper Putumayo River (B. Guevara). 

Locality interpretation. — The type locality isn't 
precise. Perhaps this can be determined by research 
on the travels of the collector. Like that of most of 
its congeners, the locality may be assumed to be 
well above 1000 m. 

Name basis. — Named after the late Dr. A.B. 
Gumey formerly of the U.S. National Museum's en- 
tomological staff who assisted my early embiid re- 
search. 

Discussion. — My revised illustration of the ho- 
lotype shows that C. gurneyi is distinct in the re- 
duced sclerotization of the epiproct (EP), fairly dis- 
tinct microspiculation of the crest of the medial flap 
(MF); the small, blunt, right tergal process (10 RP) 
tucked under the caudal margin of 10 R; the longer, 
triangular, membranous apex of the hypandrium 
process (HP), the large, quadrate, caudal extension 
of the setose lobe of the left paraproct (LPPT); the 
absence of an isolated, small sclerite between this 
lobe and the paraproct's sclerite at the left cercus 
base: the relatively short left cercus lobe, and only 



ROSS: EMBIA. BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



19 




RGB 



LP 



10 RP LPPT 

10 LP 

TERMINALIA ventral 



Figure 5. Calamoclostes albistriolatus Enderlein, topotype. Type locality: Banos, Ecuador. 



short whiteness on the few cross-veins. My only 
specimens with most of these characters are from 14 
mi SE Caqueza, Meta Prov.. Colombia, 1300 m, but 
this place is far to the north of the upper Putumayo 
type locality. 

During 1955 I also reared a large series related 
to C. giimeyi, 2 mi W of Alban, Cundin Amarca 
Prov, Colombia, 2020 m, under bark flakes of trees 
in a pasture. Males have larger wings with fewer and 
less white cross-veins; EP is well sclerotized; MF 
crest is strongly microspiculate; 10 RP is triangular, 
projected caudad; HP process is more rounded, and 
the membranous portion of LPPT is rounded, not 
quadrate. The series probably represents a new 
species worthy of a name when a comprehensive 
study of Calamoclostes of the Bogota region is 
made. 



Calamoclostes auriceps Ross 
new species 

Holotype. — Male, on slide, CAS. Data. — Ecua- 
dor: Pistishi, 8 mi S. Alausi, Chimborazo, 2400 m, 
colony in crack in vertical, silty, road bank, ma- 
tured in culture 11-III-55 (E. S. Ross and E. I. 
Schlinger). 

Name basis. — Auriceps in reference to golden 
head. 

Description. — Appearance: Apterous, \ery 
large (body length 14.0 mm); body sclerites and 
legs mottled chestnut brown, membranes cream 
white; pale sclerites and adjacent membranes form- 
ing a pale band between each thoracic somite; head 
extensively golden. Color details (in alcohol): Cra- 
nium golden except for a transxerse. chestnut 
brown diamond-shaped area between antennae and 



20 



OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES 




Figure 6. Calamodostes gurneyi Ross, holotype. Type 
locality: Upper Putumayo R.. Colombia. 



a cloud on vertex, thus delimiting a transverse gold 
band between very small, black eyes. Venter of 
head and mouthparts lemon yellow; antennae light 
tan. Dorsal and pleural scierites of thorax mottled 
chestnut brown; prosternum and cervical scierites 
chestnut brown, meso- and metathoracic sterna yel- 
low brown, irregularly margined with cream white, 
adjacent membranes cream white; acrotergites 
translucent amber, adjacent dorsal membranes 
cream white, thus forming two intersomital pale 
bands. Legs basically concolorous with thoracic 
terga except for creain white femora-tibial joints on 
hind legs. Abdominal terga 1-5 concolorous with 
thoracic terga, 6-8 increasingly pale caudad due to 
medial, cream white clouding; sterna 1-8 and pleu- 
rites golden, sternite 9 chesmut brown; pleural 
membranes cream white. Tenth tergite's hemiter- 
gites dark chestnut brown, left process piceous; 
membranes cream white. Cerci tan with joint mem- 
branes cream white; left cercus largely golden 
brown, apical half of lobe dull yellow. 



Anatomical distinctions; Cranium nymphaform, 
eyes very small. Antennae very slender, segments 
small, 24 (complete); body and legs essentially sim- 
ilar to a female. Medial papilla of hind basitarsus in- 
flated, conspicuous. Left tergal process (10 LP) pi- 
ceous, short, curved down, broad, surface rugose 
(the contact point with left cercus lobe) apex a sharp 
spine, not visible from above; right tergal process 
(10 RP) triangular, weakly sclerotized. Medial flap 
(MF) carinate, not microspiculate at apex. Epiproct 
sclerite (EP) broadly attached to MF, thence tapered 
caudad. Basal seginent of left cercus very stout, its 
lobe very large, curved basad, dorso-ventrally flat- 
tened (pinched); basal segment of right cercus stout 
on inner side, greatly expanded distad, apical seg- 
ments tapered distad, mahogany brown with ex- 
treme apex cream white; joint membranes gray. 

Allotype. — Female, in alcohol (CAS). Data: 
From holotype 's culture. 

Description. — Basically similar to male but not 
as dark. Cranium with dorsal pattern outlined in 
brown. Thoracic bands paler, inore conspicuous. 
Body length: 18.0 mm. 

Paratypes. — Males from type culture deposited 
in CAS, USNM, and Ecuador's Museo de Ciencias 
Naturales, Quito. 

Calamodostes inicropterus Ross 
new species 

(Figure 7) 

Holotype. — Male, on slide, CAS. Data. — Ecua- 
dor: 1 1 mi E. Paute, Azuay, 2800 m (est.), 17-11-55 
(E. S. Ross). 

Description. — Appearance: Rather large (body 
length 13.5 mm). Body and legs varied shades of 
light chestnut brown except prothorax and its legs, 
which are dark chestnut brown, and a pale band be- 
tween pro- and mesothorax. Color details (in alco- 
hol): Cranium very dark chestnut brown, lacking a 
golden band between eyes; area around anterior ten- 
torial pits faintly golden. Basal antennal segments 
medium brown, others grading to tan at apex, 22 
segmented (complete). Acrotergite 1, pale tan, adja- 
cent membranes gray white, thus forming a pale 
thoracic band. Sclerotic portions of terminalia var- 
ied shades of very dark chestnut brown to piceous, 
especially on medial sclerite (MS), the left tergal 
process (10 LP), and margins of basal segment of 
left cercus; basal segment of right cercus sclerotized 



ROSS: EMBIA. BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



21 



only on convex inner side; distal segments medium 
brown, tips and intersegmental joints pale tan; cleft 
membrane of tenth tergite gray white. 

Anatomical distinctions: The very small wings 
(not nymphal pads), barely extended beyond caudal 
end of each bearing somite. Left process (10 LP) of 
terminalia has an elevated dorsal projection and a 
small, left-turned apical hook. Hypandrium process 
(HP) triangulate, weakly sclerotized. Left paraproct 
(LPPT) globose, membranous except for an isolat- 
ed medial sclerite; otherwise its sclerotic portion 
appresses the inner base of the left cercus. Basal 
segment of left cercus exceptionally stout. 

Allotype. — Female, in alcohol, CAS. Data. — 
From holotypes culture. 

Description. — Appearance: Large: generally 
amber with cream membranes, prothorax darker; 
head bright chestnut brown. Color details: Cranium 
uniformly chestnut brown, without pattern except 
for a narrow, caudally-arcing, yellow area extend- 
ing from eye to eye; clypeo-frons area dark chest- 
nut brown. Basal antennal segment amber, others to 
apex gradually blending from tan to cream (in two 
subapical segments), distal segment tan. Pronotum 
dark amber. Acrotergite 1 amber, adjacent mem- 
branes cream. Mesonotum dark amber, anterior 
fifth whitish, thus enhancing the conspicuous pale 
band between prothorax and mesonotum. Metan- 
omm and all abdominal tergites pale amber with 
cream membranes. Legs varied shades of amber. 
Ventral body sclerites (except for darker proster- 
num), largely transparent cream white. Paragenital 
sclerites amber, medial areas cream. Cerci very 
pale tan, joints white. Body length; 17.5 mm. 

Paratypes. — A few topotypic adult males, CAS 
and NMQ and a few adult female parallotypes. 

Discussion. — From C. auriceps, the only other 
Calamoclostes in the same high Andean altiplano 
zone, C. micropterus is, of course, readily distin- 
guished by its miniature wings, C auriceps has 
none. However, one shouldn't be surprised if fully 
winged, or completely apterous males of this 
species occur in the same general region. Calamo- 
clostes auriceps males also differ in having a com- 
pletely golden head, that of C. micropterus is even- 
ly brown. There are numerous terminalia differenc- 
es, such as the more stubby 1 LP of C. auriceps 
and the lack of an isolated sclerite on the left para- 
proct s membranous lobe. 




TERMINALIA 
VENTRAL 



Figure 7. Calamoclostes micropterus, n. sp. holotype. 
Type locality; Near Paute, Azuay, Ecuador. 



Calamoclostes silvestris Ross 
new species 

Holotype. — Male, on slide, CAS. Data. — Ecua- 
dor: 15 mi SE Santa Rosa, El Oro, matured in cul- 
ture 11-55 from bark flakes in dark primary rainfor- 
est (E. S. Ross and E. I. Schlinger). 

Name basis. — Refers to occurrence m lowland 
rainforest (unusual for the genus). 

Description. — Appearance: Alate, very dark 
brown throughout, head darker. Color details (in 
alcohol); Cranium very dark chocolate brown with 
two very faint, transverse, diffused, oval, golden 
spots between eyes. Antennae concolorous with 
cranium, 24-segmented (complete). Palpi, mentum 
and anterior third of submentum dark brown, be- 
coming chestnut brown caudally. Pronotum medi- 
um brown, becoming pale laterally, adjacent mem- 
branes cream white. Pterothorax and legs varied 
shades of dark brown, hind basitarsus and its setae 
especially dark. Wings small for the genus with nar- 



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row, white, costal and apical margins; veins weakly 
pigmented; one cross-vein between RP and MA, 
one between RP and MA, +2- two between MA and 
MP, all white when crossing exceptionally narrow 
hyaline stripes. Abdomen, including terminalia, var- 
ied shades of brown; pleurae, especially toward 
base, cream white. 

Anatomical distinctions. — Mandibles short, 
broad; the left one with a large incisor flange. Eyes 
rather small, three eye-width interspace. Hind ba- 
sitarsal mid-papilla large. Left tergal process (10 
LP) short, down-curved, caudal surface rugose, ter- 
minated ventrally as a narrow, sharp point turned 
leftward. Forward end of medial flap (MF) densely 
microspiculate. Apex of hypandrium lobe (HP) 
membranous, triangular. Left paraproct with a sec- 
ond non-setose sclerite in membrane. Dimensions 
(on slide): Body length 11.0 mm; forewing length 
6.7 mm., breadth 1.6 mm. 

Allotype: — Female (in alcohol) CAS. Data: 
From holotype's culture. 

Description. — Appearance: Varied shades of 
brown, legs and antennae golden tan. Color details: 
Cranium uniformly dark chestnut brown with a faint 
golden chevron between black eyes. Antennae gold- 
en tan, all but seven segments lost. Mouthparts and 
ventral area golden brown. Pronotum light chestnut 
brown. Meso- and metathoracic scuta golden brown 
medially, blending to dark lavender brown on sides 
and across apices; abdominal terga similar. Ventral 
sclerites, including paraprocts and all legs, golden 
tan; pleurae cream white; cerci tan, mottled with 
brown. Body length: 10.0 mm., actually much 
longer because the specimen died and the body seg- 
ments telescoped. 

Paratype. — One male (CAS) from holotype cul- 
ture. 

Discussion. — The occurrence of a Calamo- 
clostes in dark, virgin forest at low altitude on the 
west side of the Andes, is exceptional. Males are 
distinct in having almost the entire costal and apical 
margins of the wings narrowly white, all congeners 
have brown margins. The terminalia are much like 
those of C. albistriolatus except for the stubby, ex- 
panded, rugose, abruptly downwardly-hooked apex 
of 10 LP (apex scarcely visible from above); the 
more acute apex of HP; the presence of a detached, 
non-setose sclerite just mesad of the sclerotic por- 



tion of LPPT, and the more expanded, pale apex of 
the left cercus lobe. 

Calamoclostes oculeus Ross 
new species 

Holotype. — Male, on slide, CAS. Data. — Co- 
lombia: Salado, 27 mi. W. of Cali, 1350 m, 21-III- 
55 (E. S. Ross, E. L Schlinger). Under bark flakes 
of shade trees in plaza. 

Name basis. — Latin oculeus, full of eyes, in ref- 
erence to the relatively large eyes of the species. 

Description. — Appearance: Alate, rather small 
for the genus, uniformly brown; head darker, lack- 
ing maculation. Color details: Antennae concolor- 
ous with cranium, as are the mouthparts. Prothorax 
slightly paler than pterothorax. Pterothorax and legs 
varied shades of medium brown, hind basitarsus not 
especially dark. Wing bands light brown, hyaline 
stripes rather broad, MA forked beyond apical half 
(not forked in some specimens), cross-veins absent 
behind RP. Abdomen, including terminalia, concol- 
orous with pterothorax. 

Anatomical distinctions. — Cranium somewhat 
quadrate. Eyes especially large, inflated; half of 
head length long, separated by slightly more than an 
eye width. Mid-papilla of hind basitarsus perhaps 
absent, at least not inflated or readily discernable. 
Terminalia basically as in C. albistriolatus but less 
sclerotic, but its 10 LP has a shorter sub-basal spur 
on its left side and a tiny one opposite on the right 
base; HP margin broadly rounded, membranous; in- 
ner setose portion of LPPT large, globular. Dimen- 
sions (on slide): Body length 9.5 mm; forewing 
length 8.0 mm, breadth 2.0 mm. 

Allotype. — Female, in alcohol, CAS. Data. — 
From holotype's culture. 

Description. — Appearance: Smallest species of 
the genus (body length 10.0 mm); body and legs 
uniformly tan except for a slightly darker prothorax 
and legs; all body membranes pale tan, almost same 
tone as sclerites, no pale thoracic intersomital 
bands. Cranium uniformly chestnut brown except 
for two, suffused golden areas between eyes. Anten- 
nal segments 1-15 uniformly light brown, 16 and 17 
golden tan, 18 light brown (segmentation com- 
plete). Paragenital sternites medium brown. Para- 
procts translucent pale tan. Basal segments of cerci 
pale golden tan, distal segments light golden brown. 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



23 



Paratypes and parallotypes. — Numerous topo- 
typic males and females deposited in CAS, USNM, 
BMNH, and Institute de Historia Naturales, Bogo- 
ta, Colombia. 

Microembiinae Ross 

new subfamily 

Type genus. — Microembia Ross, 1 944, by pres- 
ent designation. 

Distribution. — South America: Upper Amazon 
(one record). 

Diagnosis. — Male (Fig. 8): Very small (body 
length about 5 mm), alate, pale tan throughout. 
Head circular, dorsoventrally thick. Eyes large, 
wide-spaced, extending caudad half of the cranium's 
length, facets very large; clypeus and frons heavily 
sclerotized rugose, anterior angles strongly lobed 
forward and dorsad. Mandibles thin, relatively large 
due to great expansion of outer basal angles; distal 
teeth small. Submentum well sclerotized; anterior 
margin transverse, weak; lateral margins strongly 
inflexed, convergent caudad. Antennal segments ex- 
ceptionally long, segments beyond the fifth broken 
off. Wings narrowly tapered basad, very pale due to 
reduced venal bands and broad hyaline intervenal 
stripes; venation embioid with all veins unsclero- 
tized except for blood sinuses, RP and RP + MA; 
cross-veins absent except for one between apex of 
RBS and RP. Legs very long and slender; hind ba- 
sitarsus lacking a medial papilla. Terminalia small, 
weakly sclerotized. Ninth tergite (9) membranous 
except for basal margin and outer caudal angles. 
Basal margin of tergite 10 narrowly sclerotized, 
slightly arcuated basad; the tergite broadly cleft to 
base; left hemitergite ( 10 L) well sclerotized except 
at its weak inner base, caudal margin arcuately pro- 
jected caudad; inner margin straight, continuous 
with that of its process ( 10 LP) which is narrow, par- 
allel-sided and simple; inner margin of right 
hemitergite (10 R) weak, blending with membrane 
of tergal cleft, outer margin strong, evenly acuated 
from outer base to caudal apex; right process (10 
RP) a short, blunt, fleshy lobe folded beneath 
caudo-mesal margin of 10 R. An obscure sclerite on 
the inner margin of 10 R may be the homolog of a 
median flap (MF?); beneath it a caudally-rounded 
lobe may be the epiproct (HP?). Ninth sternite (H) 
broad basally, evenly tapered caudad, becoming an 
angular process (HP). Left paraproct (LPPT) very 



large, closely contacting H and HP on inner side; 
right paraproct (RPPT) obsolete, with only weakly 
sclerotized fragments. Cerci short, left cercus two- 
segmented, apical segment broadly attached to the 
basal segment (LC,) which is well sclerotized, dis- 
tally enlarged, but not abruptly lobed, inner margin 
of this expansion coarsely echinulated; right cercus 
normally articulated; both cerci, especially ventral- 
ly, bearing numerous trichobothria with large sock- 
ets. 

Females. — Unknown. 

Discussion. — This category is proposed with 
hesitation because of the limited series and rather 
indecisive characters. The subfamily seems to be 
most closely related to Archembiinae but the com- 
ponent species differs in its minute size, reduced 
wing vein sclerotization, lack of a second basitarsal 
papilla, the terminalia s lack of a well-developed 
medial flap, and reduced lobing of the left cercus. 
Judging from its large eyes and pale coloration, 
males of the type species must disperse nocturnally. 
This is unusual for an embiid inhabiting tropical 
forests. 

The subfamily contains the one genus, Mi- 
croembia Ross (1944:416) and species, M. rii- 
gosifrons Ross, I.e. (holotype male, on slide 
USNM; one additional male in my collection, CAS) 
(Fig. 8). 

These, the only specimens collected to date, 
were secured at Iquitos, Peru, March- April 1931 by 
R. C. Shannon. They were probably attracted to 
light. During a visit to Iquitos I atternpted without 
success to locate colonies of the species. These must 
be very obscure in bark crevices, but may no longer 
occur within the limits of the now much-disturbed 
city. On remnant trees along the river front 1 found 
only numerous colonies of the "weed species," 
Archembia batesi. 

Szumik's suggestion (1996) that Microembia 
has a relationship with Ptiloceremhia of SE Asia, 
based in part on the highly unreliable group charac- 
ter — the absence of a middle basitarsal papilla, is 
weak. Her discussion of the relationship to other 
embiids is inconclusive. I am inclined to relate the 
genus to the Archembiinae. Until more fieldwork in 
the Amazon Basin yields related species or genera, 
Microembia will "hang in taxonomic space." 

Although not used in the present systematic 
studies, 1 have accurately included chaetotaxy in m\' 
revised illustration of the terminalia w hich is based 



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SUBMENTUM 



TERMINALIA 

VENTRAL 
Figure 8. Microemhia nigosifrons Ross, paratype. Type locality: Iquitos. Peru. 



on the CAS paratype. Microembia has an unusually 
large number of large-socketed trichobothria on the 
cerci of males. 

Scelembiinae Ross 

new subfamil} 

Type genus. Scelembia Ross. 1960, by present 
designation. 



Distribution. — Afrotropical and Neotropical re- 



eions. 



Diagnosis. — Males small to moderately large 
(body length 6-18 mm), usually alate: pale tan to 
black, often with orange or reddish prothorax and/or 



pterothorax. Cranium usually elongate-oval; eyes 
small to moderately large: clypeus and frons not es- 
pecially sclerotic or rugose, anterior angles not pro- 
duced; mandibles dorso-ventrally thin, apical teeth 
well spaced on a horizontal plane (not curved ven- 
trad), mandibles not dentate in some African gen- 
era; submentum usually weakly sclerotized with 
apical and lateral margins not inflexed (except in 
three Neotropical genera, Ischnembia. Pararha- 
gadochir and Litosembia, as well as in several 
African genera). Wings highly variable in size and 
pigmentation, absent in many species (all species 
of some African genera); MA usually forked. Ninth 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



25 



abdominal tergite extensively membranous; tergite 
10 broadly cleft to base. Medial sclerite (MS) form- 
ing the narrow, transverse basal margin of tergite 10 
(in some Afi^can genera an often detached, isolated 
sclerite, present in the cleft membrane, may be MS). 
Medial flap (MF) usually a diagonal sclerotized arm 
projected forward in cleft (never a flap, as in Arch- 
embinae); in some American genera MF is obsolete, 
represented only by membranous wrinkles. Left 
paraproct (LPPT) well developed, at times bearing a 
micro-echinulate nodule, or an acute point on inner 
caudal angle, but never a distinct process. Basal seg- 
ment of left cercus usually lobed medially or distal- 
ly, these lobes always echinulate; however, in 
Mesoamerican and some Mexican genera (except 
for introduced Pararhagadochir) the lobe is at ex- 
treme base of the segment and may not be echinu- 
late. Conocercembia of Mexico has a prominent 
distal lobe, but this isn't echinulate. Apical cercus 
segments always present, well articulated. 

Females. — Not studied for subfamily characters. 

Discussion. — As evident in the above descrip- 
tion, this large, widespread subfamily is highly di- 
versified and difficult to define. However, especial- 
ly in the New World, an experienced specialist can 
readily recognize its genera. The subfamily's great- 
est diversity occurs in Africa, the locale of its type 
genus, Scelembia Ross. Except for the teratembiid 
genus Diradius Friederichs, it is the only embiid 
group with strong transatlantic relationships. 

KEY TO NEW WORLD GENERA OF 
SCELEMBIINAE 

(Adult males) 

1. Tertiary shale fossil, Florissant Colorado 
Lithembia 

— Extant species 2 

2. Left cercus lobe located medially or distally, al- 
ways echinulate. Endemic to South America 
3 

— Left cercus lobe confined to extreme base, at 
times not echinulate. If a distal lobe is present, 
it is not echinulate (except for introduced 
Pararhagadochir in which it is echinulate). En- 
demic to Mexico and Central America 14 

3. Membrane at forward end of MF not mi- 
crospiculate in a depression 4 

— Membranes at forward end of MF depressed 
and microspiculate 12 

4. Medial flap of tenth abdominal tergite (MF) 



vestigial, represented only by membranous 
wrinkles 5 

— Medial flap conspicuous, an at least partly 
sclerotic arm projected forward in cleft mem- 
brane 7 

5. Left tergal process (10 LP) simple - a large, 
sclerotic talon arcing leftward Ochrembia 

— Left tergal process bifid, consisting of a small 
inner talon and an outer, often submembranous 
flange 6 

6. Surface of left hemitergite (10 L) diagonally 
ridged, continued meso-caudad as the base of 
the inner talon of its process (10 LP); outer 
flange broad extended partly beneath the talon 
Ambonembia 

— Surface of 10 L almost flat; talon and flange of 
left tergal process, small, widely separated 
Mahjcosembia 

7. Process of left hemitergite simple, consisting 
only of an acutely tapered talon 8 

— Process of left hemitergite bifid 11 

8. Base of left tergal process (10 LP) arced for- 
ward almost contacting base of tenth tergite. 
Caudal margin of left hemitergite (10 L) ob- 
tusely angulate Dolonembia 

— Base of 1 LP not arced forward. Caudal mar- 
gin of 10 L straight or tapered caudad 9 

9. Cerci almost entirely white (not melanized), 
right cercus entirely white. Small species 
Argocercembia 

— All segments of cerci brown (melanized). 
Small to large species 10 

10. Very large bicolorous species. Caudal margin 
of left hemitergite ( 10 L) membranous; medial 
flap (MF) elongate, narrow; talon of right pro- 
cess (10 RP) small, twisted toward right. At- 
lantic forests of east-central Brazil 

Embolyntha 

— Small unicolorous species. Caudal margin of 
10 L not membranous. Medial flap broad, 
short; talon of right process large, angled 
mesad. Amazon Basin Xiphosembia 

11. Process of left hemitergite with talon and 
flange almost equal in length, and well spaced 
Bigiiembia 

— Talon of left process much larger than flange, 
the latter at times partly hidden beneath base 
of talon Gibocercus 

12. Very small, pale, desclerotized males. Medial 
flap (MF) terminating at forward end as a 
small, but distinct, microspiculate nodule. Tal- 



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OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES 



on of left tergal process minute, not separated 
from the broad, faintly-visible flange. Belem, 
Brazil Litosembia 

— Small to very large, usually well sclerotized. 
Medial flap's forward end with a concave, mi- 
crospiculate area in cleft membrane. Talon and 
flange of left process well developed and well 
separated 13 

13. Small, pale males. Concave microspiculate area 
in cleft membrane very small. Inner basal side 

of left cercus deeply depressed 

Ischnosembia 

— Medium to large males. Concave, microspicu- 
late area in cleft membrane large, conspicuous. 
Inner basal side of left cercus not depressed 
Pararhagadochir 

14. Lobe at inner base of left cercus echinulate 
Neorhagadochir 

— Base of left cercus not distinctly lobed or 
echinulate 15 

15. Left cercus with a distal, acute, non-echinulate 
inner lobe Conocercembia 

— Left cercus without a distal inner lobe 
Brachypterembia 

Genus Lithembia Ross 

Lithe?nbia Ross, 1984:83.— Carpenter, 1992:190. 

Type species. — Embia florissantensis Cockerell. 

Distribution. — Oligocene fossil in volcanic ash 
shale, Florissant, Colorado (one specimen). 

Discussion. — The type is a large male with fair- 
ly clear wings but only a dark blotch in the position 
of the male terminalia. The wings resemble those of 
the family Embiidae, vein MA being clearly forked. 
No details of the terminalia can be ascertained. In 
view of the fossil's large size, it is not likely to be a 
teratembiid and the forked MA rules out assignment 
to Anisembiidae. Its slender body and northerly oc- 
currence indicates that it isn't a clothodid, a family 
which appears to ha\'e always been restricted to 
South .America with an extension into Panama. 
Therefore, like all other North and Central Ameri- 
can embiids with MA forked, it must be placed in 
Embiidae. more precisely in Scelembiinae. 



Lithembia florissantensis (Cockerell) 

Embia florissantensis Cockerell, 1908:230, fig. 4. — Han- 
dlirsch. 1906-08:1357.— Enderlein, 1912:53. 

Oligotoma florissantensis (Cockerell), Krauss. 1911:48. 

Clothoda florissantensis (Cockerell). Davis, 1939:379. — 
Ross, 1944:406. 

Lithembia florissantensis (Cockerell), Ross, 1984:83. — 
Carpenter, 1992:190. 

Holotype. — Alate male on rock slab in Riker 
Mount, Univ. of Colorado Museum, Denver. 
Data.— Florissant Colorado Station 14, 1907[?] 
(W. P. Cockerell). Miocene. 

Discussion. — All references are based on Cock- 
erell's original publication. No additional speci- 
mens have been found. The writer has studied the 
type and cannot add new details to the description, 
or improve on the original published photograph, 
except to note that it is reversed, perhaps due to an 
inverted negative. 

Even if additional specimens are collected, it is 
doubtful if they would reveal sufficient terminalia 
details to add significant information concerning 
the relationships of the species. 

The occurrence of an embiid in the ancient 
Florissant Basin, about thirty-five million years 
ago. indicates that the climate at that time lacked a 
prolonged cold winter. At the time of fossilization 
the area is reported to have had an estimated eleva- 
tion of only 900 meters. Therefore, the estimate of 
about 2000 meters by K.M. Gregory would appear 
to have been too high for embiid survival at Floris- 
sant's latitude. Of course, one must consider in- 
crease in altimde and changed geographic position 
as a result of orogeny and tectonic movement. 

Genus Embolyntha Davis 

Embohmtha Davis, 1940b;344 (new name for Olyntha 
Gray,. 1832). Ross, 1944:412. 

Olyntha Gray. 1832:347.— Burmeister, 1839:70.— Davis, 
I.e. (as a name preoccupied by Olynthus Hiibner, 1819). 

Embius Gray,1832:786:72, fig. 2 (as a name preoccupied 
by Embia Latreille. 1829). — Davis, I.e. 

Type species. — Olyntha brasiliensis Gray, 
1832, by original designation. 

Distribution. — Southeastern Brazil. 

Diagnosis. — Males: Very large (body length 
more than 18 mm), robust; strongly pigmented. 



ROSS: EMBIA. BIOSYSTEMATICS OF THE ORDER EMBIIDINA. PART 3 



27 



blackish brown with prothorax. coxae, trochanters 
and femora of fore- and niidlegs bright orange, an- 
tennal apices and numerous cross-veins of wings 
white. Cranium elongate-oval with pronounced lon- 
gitudinal sulci paralleling lateral margins. Eyes 
small. Antennae very long (30-segmented, com- 
plete), segments short, setae fine, segments 1-24 
uniformly dark, apical six pure white. Mandibles 
delicate, short, outer margins evenly arcuate; apical 
teeth small, even in spacing and size. Mentum con- 
spicuous; submentum quadrate, evenly but not 
strongly sclerotized. Wings very large; vannal area 
reduced, narrowly tapered; all veins strong with 
considerable variation in branching and in the num- 
ber and position of cross-veins (even in a single 
specimen); at times MA branches and MP are sec- 
ondarily branched. Hind basitarsus densely setose, 
medial papilla small. Abdominal terminalia highly 
asymmetrical due to great size of left hemitergite 
(10 L). Ninth tergite very narrow on left side due to 
basal projection of 10 L. Left hemitergite (10 L) 
broader than long, basal and mesal margins form- 
ing a continuous sclerotic arc; caudal margin 
straight, transverse, sclerotic but with its edge 
membranous and setose. Left process (10 LP) con- 
tinuous with inner arc of 10 L, long, tapering to an 
acute point on outer side with a subterminal trun- 
cate flange interrupting inner side. Medial cleft 
very narrow, paralleling basal and medial arc of 1 
L and its process. Right hemitergite (10 R) termi- 
nated as a narrow, ventrally projected, sinuous, 
spine Jike process (10 RP). Medial flap (MF) nar- 
row, sclerotized ventrally only. Epiproct (EP) a very 
narrow fleshy lobe as long as 1 LP, bearing a very 
slender diagonal sclerite which extends beneath 1 
LP to inner base of left cercus. Ninth sternite (H) 
very broad; its process (HP), which is not projected 
much beyond caudal margin of H, is simply a scle- 
rotic fold. Left paraproct (LPPT) small, inner-cau- 
dal angle bearing a small nodule terminated by a 
narrow, microspiculate appendix. Basal segment of 
left cercus very short, sclerotic; inner lobe medial, 
its basal surface extensively depressed, almost 
causing bilobing of its coarsely echinulate apex (the 
greatly produced epiproct, stiffened by its long 
sclerite, together with 10 LP, apparently fits into 
this socket). 

Females: Very large (body length averaging 25 
mm), unicolorous blackish brown including mem- 
branous areas; legs and venter of thorax mostly 



golden. Antennal segments 31-33, brown, distal 
segments contrastingly white. Cerci uniformly dark 
brown. Hind basitarsi with two large ventral papil- 
lae. Eighth sternite strongly lobed medio-caudally, 
sides darkly pigmented; first valvifer lobes promi- 
nent; second valvifer represented by clear amber, 
lateral plates. Nmth sternite uniformly sclerotized, 
dark brown. 

Discussion. — At this time Embolyntha is limit- 
ed to its very distinct type species which is readily 
identified by generic-level characters. Davis 
(I940b:345) redescribed the holotype, an adult male 
in the British Museum (N.H.) with only "Brasil" as 
locality data. Recent collecting suggests that this 
type must have been collected in, or near. Rio de 
Janeiro. 

Although it is one of the first named species of 
the order, one having exceptionally large body size, 
and occurs in the vicinity of one of the worlds great 
cities, specimens of E. brasiliensis are very rare in 
collections. Perhaps less than six adult males have 
been collected to date and females and the habitat 
have remained unknown. However, after much 
search, I finally discovered that the species incon- 
spicuously inhabits mossy rock ledges and road 
banks in rainforest near Paineiras in the heights 
above Rio de Janeiro. Each young nymph was de- 
veloping alone in a separate gallery obscured in 
moss and other growth. The galleries extended from 
a crevice retreat into surface moss. After much ef- 
fort, 1 collected and cultured ten nymphs but, unfor- 
tunately, all matured as females. It must not be as- 
sumed that the species is at times parthenogenetic, 
for the unmated females laid infertile eggs. In addi- 
tion to these females, my collection (CAS) contains 
an adult male from Brazils Parque National do Ita- 
tiaia, 26-X-58 (D. Lacombe) caught in diurnal flight 
near a mossy road bank. 

It is probable that the brood of each female soon 
disperses into scattered crevices and thus the spe- 
cies may never produce the conspicuous, sheet-like 
colonies characteristic of the most common Rio de 
Janeiro embiids (Archemhia spp.) found in the same 
habitat. 

Embolyntha brasiliensis (Gray) 

(FldURE '^) 

Emhiiis (?) brasiliensis Gray, 1832:786. pi. 72. fig. 2. 

Olyntha hriisiliensis (Gray), Gray. 1832:347. — West- 
wood, 1837:373, pi. 2. fig. 3. — Burmeister. 



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OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES 



1839:770.— Walker, 1853:532. — Krauss. 1911:28, pi. 
l,fig. 1. 

Embia {Olyntha) brasiliensis (Gray), Hagen, 1885:195. 

Embia brasiliensis (Gray), Enderlein, 1912:48, fig. 24. — 
Navas, 1918:98,— Costa Lima, 1938:109, figs. 52-54. 

Embolyntha brasiliensis (Gray), Davis, 1940b:345, figs. 
1-7 (redesc. Type).— Ross, 1944:413. 

Embia brasiliensis \ax. flavicercatus Enderlein, 1912:49 
(prob. a distinct sp.). 

Type.— Male, Children Collection, BMNH. 
Type labels. — "Brasiliensis G.R. Gray Brasil,"" "40 
3.30 704," "Type" (red-circled card). 

Condition. — Pinned dry with wings spread, 
mouthparts dissected on card, genitalia preserved in 
small vial attached to pin. General condition excel- 
lent. 

Remarks. — My examination of the type con- 
firms the accuracy of current identifications. Be- 
cause most specimens were collected in or near Rio 
de Janeiro, the type locality is here formally estab- 
lished as Paineiras, 450 m, Parque Nac. do Tijuca, a 
well protected environment. The present generic 
treatment and figure, are based on a male (CAS) 
from Parque Nacional do Itatiaia, Est. do Rio de 
Janeiro. 

The varietal name Jlavicei-cata Enderlein, prob- 
ably pertains to a distinct species — perhaps of the 
genus Archemhia. Enderlein apparently had only a 
literature knowledge of a Burmeister specimen. Be- 
cause no locality is specified and the specimen may 
not be preserved, the varietal name need not be rec- 
ognized, if indeed it ever had nomenclatural status. 

I attribute the name E. brasiliensis to Gray, 
rather than Griffith and Pidgeon, as was done by 
Davis (1940b), because Gray authored the portion 
of "Animal Kingdom" dealing with Embiidina. 

Specimens examined. — 1 male (Copenhagen 
Museum) Brazil: "Rio Reinh." (I assume that "Rio" 
is the locality and "Reinh.," an abbreviation of the 
collector's name. 1 male (USNM) Rio de Janeiro, 
Yellow Fever Survey (R.C. Shannon); 1 male (CAS) 
Itatiaia, Est. do Rio 26.X.58 (D. Lacombe); 8 adult 
females (CAS), Paineiras, Parque Nac. do Tijuca. 
each died infertile during VIII and IX-64 and III-65 
(E. S. Ross). 

Perhaps because of rarity, E. brasiliensis has no 
synonyms. Costa Lima's treatment (1938:109-1 14), 
as evidenced by his figures 53 and 54, was based on 
correctly identified specimens of £. brasiliensis. All 



of the anatomical work of Lacombe and Barth, 
using the name brasiliensis, was based on speci- 
mens of Archembia lacombea Ross. 

Genus Xiphosembia Ross 

new genus 

Type species. — Xiphosembia amapae Ross, new 
species by present designation. 

Distribution. — South America: Amazon Basin. 

Name basis. — Greek xiphos (sword or saber) in 
reference to the saber-like left tergal process of the 
terminalia. 

Diagnosis. — Males: Medium sized (body length 
8-12 mm), alate; pale to darkly pigmented. Crani- 
um variable in form; eyes small to large; antennae 
without pale distal segments, 22 to 28-segmented; 
submentum transversely rectangulate, anterior mar- 
gin weak; mandibles flat, thin, short, apical teeth 
well spaced and acute. Hind basitarsus with two 
ventral papillae. Tenth tergite very broadly cleft to 
basal margin. Left tergal process (10 LP) very large, 
arising on inner side of the hemitergite; arcuate, nar- 
row, evenly-tapered to a fine point; lacking an outer, 
basal appendix or flange except for a tiny, obscure 
one in a new species from Rondonia. Right hemiter- 
gite deeply excised on inner-basal side; process (10 
RP) an abruptly-narrowed meso-ventrally-directed, 
large talon. Medial flap (MP) a wrinkled, sclero- 
tized area directed meso-caudad, without microspic- 
ulations at forward end. Epiproct (EP) large; its 
sclerite narrow, inconspicuous. Hypandrium 
process (HP) truncate, not attaining caudal margin 
of left paraproct. Left paraproct (LPPT) with caudal 
margin simple, microspiculate in type species, but 
sclerotic and bilobed in another species. Basal seg- 
ment of left cercus with a single, large, echinulate, 
inner-apical lobe; outer side well sclerotized. 

Females: Without noteworthy integumental 
characters. Coloration very distinctive: antennae 
white-tipped; prothorax and mesothorax and legs 
yellow in all species; abdomen dark brown in one 
species, yellow in two species; head dark brown in 
all species. 

Discussion. — The simple, scimitar-like left ter- 
gal process in combination with a prominent medi- 
al flap are distinct features of the male terminalia. 
Three species of Xiphosembia are represented in my 
collection. One from south of Belem is very closely 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, R\RT 3 



29 




TERMINALIA 

VENTRAL 

Figure 9. Embolyntha brasiliensis (Gray). Type locality: Assumed to be Rio de Janeiro, Brazil. 



related to X. amapae, differing primarily in its more 
elongate head with small eyes. Both have distinctly 
pigmented females — these being entirely yellow ex- 
cept for a brown head and a largely dark brown 
metanotum. 

Xiphosembia amapae Ross 
new species 

(Figure 10) 

Holotype. — Male, one slide, MNRJ. Data. — 
Brazil: Vila Amazonas, port of Icomi Mine, near 
Macapa, Amapa. Matured in culture 28-V1I-1964 
(E. S. Ross). 



Description. — Appearance: Medium sized, 
alate; pale tan with dark brown head, light brown 
terminalia and wings; antennae almost unicolorous. 
Color details (in alcohol): Cranium mottled ma- 
hogany brown with distinct vertex pattern. Eyes 
gray-black with pale outline. Antennae uniformly 
medium brown, segments 2-3 slightly paler: 22- 
segmented. Mouthparts medium brown. Pronotum 
light brown, other thoracic sclerites pale yellow 
with darker sutures: all thoracic membranes cream 
white. Fore and mid coxae, trochanter and femora 
cream white; tibiae and tarsi light brown. Hind 
coxae light brown, trochanters and femora bases 



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OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES 



cream white; femora grading to light brown distad, 
tibiae and tarsi hght brown. Wings with all veins 
sclerotized except apices of MA, MP and entire 
CU|a; cross-veins present between all veins except 
within MA fork, and behind MP; hyaline stripes 
rather broad, venal stripes light brown. Abdominal 
sclerites light brown except for darker terminalia, 
all membranes pale cream. Dimensions (on slide): 
Body length 8.5 mm; forewing length 6.0 mm, 
breadth 1 .5 mm. 

Important integumental characters. — The short 
cranium with strongly caudally-convergent sides 
which are shorter than an eye-length; the very large, 
inflated eyes which are separated by less than an 
eye-width; and the relatively short antennae with 
only 22 segments. The terminalia are distinctive in 
the mesal origin of the left tergal process (10 LP), 
which has a simple, tapered, arcuate form; the trian- 
gulate shape and striation of the medial flap (MF); 
the simple, submembranous, microspiculate caudal 
margin of the left paraproct (LPPT) and the peculiar 
form of the basal segment of the left cercus, the lobe 
being extremely long and narrow, almost as long as 
the segment, and its caudal side is submembranous. 

Allotype. — Female, in alcohol, with holotype 
data and disposition. 

Description. — Appearance; Rather small; en- 
tirely yellow except for white-tipped antennae, dark 
brown head and portions of metanotum. Color de- 
tails; Cranium dark mahogany brown, paler anteri- 
orly and ventrally. Eyes black. Basal third of anten- 
nae pale yellow blending to dark mahogany brown 
in medial area; distal four segments pure white (24- 
segmented). Mandibles and submentum dark 
amber, other sclerotized portions of mouthparts pale 
yellow. Dorsal body sclerites pale yellow except for 
metanotum which is clouded with dark mahogany 
brown in lateral third and yellow medially; meta- 
pleurae also dark mahogany; ventral sclerites and all 
membranes cream white. All legs entirely pale yel- 
low except for largely medium brown, hind legs. 
Paragenital sternites translucent pale yellow. Body 
length 9.0 mm. 

Paratypes and parallotj'pes. — Numerous adults 
from t)'pe culmre deposited in major entomological 
collections, including CAS, USNM, BMNH, MNRJ 
and MZUSR 

Biology. — This species was discovered in a rem- 
nant patch of original forest in a region subject to a 



long dry season. Situated but a few feet above river 
level, it was ground-water dependent and thus 
trunks and branches had few of the epiphytes char- 
acterizing a rainforest. Colonies are conspicuous 
chalk white patches fully exposed on the bark of 
large trees. Largest, abandoned colonies averaged 6 
<r^ 12 inches in dimension and were distributed high 
up on the trunks. During mid-March each newly 
formed colony consisted of a single parent female 
and a first or second instar brood. Adults of these 
broods began maturing late in July and their num- 
bers peaked during August and September. A close- 
ly related new species, discovered south of Belem, 
has similar habits. Another new species from the 
Arequemes region of Rondonia has an extremely 
small, subventral, outer appendix at the base of the 
left tergal process. These will be named and de- 
scribed in a future publication. 

Genus Dolonembia Ross 

new genus 

Type species - Dolonembia tapirape Ross, new 
species by present designation. 

Distribution. — Brazil: Rio Tapirape region, 
Mato Grosso (one record). 

Name basis. — Greek dolon (dagger, stiletto, 
pike) in reference to dagger-like left tergal process 
of terminalia. 

Diagnosis. — Males distinguished from distantly 
related genus, Xiphosembia, by larger size, darker 
coloration; broader mandibles; and very distinct ter- 
minalia, as follows: caudal margin of 1 L obtusely 
angulate; 10 LP originates at extreme inner-base of 
1 L and is straight instead of arcuate; MF larger 
and more extensively projected basomesad; sclerot- 
ic, bilobed caudal margin of LPPT, and more elon- 
gate and sclerotic basal segment of left cercus with 
larger and flattened lobe. Females, instead of being 
almost entirely yellow, as in Xiphosembia, are 
brown with only the pro- and metathorax yellow. 

Dolonembia tapirape Ross 
new species 

(Figure 11) 

Holotype. — Male, on slide, CAS. Data. — Bra- 
zil; Barra do Tapirape, Mato Grosso, 7-XI-64 
(Borys Malkin). 

Name basis. — Named for the Indian village 
Tapirape. Tapirape is an Indian village at the con- 
fluence of the Tapirape and Arquais Rivers. 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



31 




HIND BASITARSUS 



TERMINALIA 

VENTRAL 
Figure 10. Xiphosembia amapae Ross, n. sp., holotype. Type locality: near Macapa, Brazil. 



Description. — Appearance: Rather large, alate; 
dark mahogany brown except for golden prothorax 
and hind femora. Color details (in alcohol): Crani- 
um uniformly dark mahogany brown, lacking paler 
clouding or pattern. Eyes black. Antennae entirely 
mahogany brown, 28-segmented. Submentum 
translucent dark amber, other mouthparts ma- 
hogany brown. Cervical sclerites and those of pro- 
thorax, including acrotergite 1 transparent gold; 
membranes cream white. Pterothoracic sclerites 
light mahogany brown, membranes cream. Forelegs 
almost entirely light mahogany brown; mid- and 
hind legs similar except for yellow femora. Abdo- 



men rust mahogany brown except for dark ma- 
hogany brown terminalia, including processes and 
cerci. Dimensions (on slide): Body length 1 1.3 mm; 
forewing length 7.0 mm, breadth 1.6 mm. 

Important integumental characters. — The broad 
oval cranium with sides equal to three eye-lengths; 
eyes small, separated by almost four eye-widths; the 
long 28-segmented antennae, and the broad, short 
mandibles. The terminalia are unique in the basal 
origin of the simple, dagger-like, left tergal process 
(10 LP) which abruptly curves caudal ly thence be- 
comes long, straight and finely-tapered. Medial flap 
(MF) exceptionally large and long, extending al- 



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most to basal margin of cleft which lacks a mi- 
crospiculated depression. Right process (10 RP) 
simple, abruptly narrowed, angled mesad, apically 
blunt. Left paraproct (LPPT) narrow with caudal 
margin sclerotic, non-spiculate and bilobed on 
inner-caudal angle. Basal segment of left cercus 
broadly lobed, the lobe sclerotic on all surfaces and 
dorsally depressed. 

Allotype. — Female, in alcohol, with holotype 
data and disposition. 

Description. — Appearance: Rather large; anteri- 
or half of body and legs bright yellow except for 
dark cranium, posterior half of body very dark 
brown. Color details: Cranium piceous brown faint- 
ly clouded with mahogany brown medially, but 
without definite pattern; ventro-medially golden 
brown. Eyes black. Basal two-thirds of antennae 
uniformly dark brown, apical third (segments 20- 
28) white. Submentum and entire labium amber yel- 
low, mouthparts otherwise shades of brown. Cervi- 
cal, prothoracic and mesothoracic sclerites translu- 
cent yellow; all membranes cream white. Metatho- 
racic sclerites dark mahogany brown, sternites 
lighter brown; all membranes cream white. All 
coxae, trochanters, and femora golden yellow; tibi- 
ae and tarsi largely light mahogany brown; hind ba- 
sitarsi with two prominent ventral papillae. Dorsum 
of abdomen basally and distally dark mahogany, be- 
coming golden brown medially; sternites 1-7 trans- 
parent light brown, lateral thirds of 8 and most of 9 
dark mahogany. Cerci dark brown except at extreme 
bases. Body length 13.5 mm. 

Paratypes and parallotypes. — Small series of 
topotypic adults deposited in CAS, MNRJ, MZUSR 

Biology: The type series was collected in colo- 
nies on bark of trees growing on a forest-covered 
rocky hill. Adult males were present during January, 
February and November. 

Genus Ischnosembia Ross 

new genus 

Type species. — Ischnosembia amazonica Ross, 
new species, by original designation. 

Name basis. — Greek ischnos (withered, thin, 
weak) in reference to the frail, thin body form. 

Distribution. — South America. — Widespread in 
forested regions. 

Diagnosis. — Males: Small (body length averag- 



ing 9 mm) slender, alate; unicolorous tan, antennal 
apices and cerci white. Cranium small, eyes moder- 
ately large; antennae 1 9-segmented, segments espe- 
cially elongate with rather long and erect setae, 3 
apical segments white; mandibles small, delicate, 
oligotomoid; submentum sclerotic, shield-like, an- 
terior and lateral margins inflexed. Wings with all 
veins behind RP reduced to rows of setae, cross- 
veins usually absent behind RP. Hind basitarsus 
elongate, with only one ventral papilla. Terminalia 
with tenth tergite broadly cleft to base. Left 
hemitergite ( 10 L) with weak inner and caudal mar- 
gins; dorsal surface pinched by two transverse, in- 
curved depressions. Process (10 LP) short, broad; 
inner talon very slender, sharply accuminated, its 
even arcuation forming a semicircle which extends 
leftward across apex of a broad, desclerotized outer 
flange. Inner-basal and outer-basal margins of right 
hemitergite ( 1 R) weak; process-base short. Right 
process (10 RP) composed of a short, sclerotic, 
meso-ventrally-directed talon and a soft, ventral 
lobe. Medial flap (MF) a thin, elevated ridge with a 
small, micro-dimpled area in the cleft membrane at 
its forward end. Epiproct (EP) large, well devel- 
oped but weakly sclerotized. Left paraproct (LPPT) 
large, caudal margin straight, membranous, finely 
microspiculated especially on its rounded, outer 
comer. Basal segment of left cercus short, descle- 
rotized except at base; lobe globose, coarsely-ech- 
inulated, angled caudo-mesad; other cercus seg- 
ments not sclerotized, white. 

Females: Slender, head exceptionally small, an- 
termae with three apical segments white; body and 
legs uniformly glossy, red mahogany except for a 
conspicuous, white zone between each thoracic 
somite; cerci pale, partially white. Hind basitarsus 
with two ventral papillae. 

Discussion: — Although this genus has many 
characters similar to those of Pararhagadochir, 
there are enough distinctions to warrant separate 
generic status. This conclusion is strengthened by 
the fact that there are three known, widespread 
species having very similar characters. The absence 
of a second hind basitarsal papilla in males, and the 
presence of two in females is unusual — almost 
unique in the order. 

From those of Pararhagadochir, Ischnosembia 
males can be distinguished by their white cerci and 
white-tipped antennae; the reduced sclerotization 
of most wing-veins, usually absent cross-veins be- 



ROSS; EMBIA. BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



33 




Figure 1 1. Dolonembia lapirape Ross, n. sp., holotype. Type locality: Barra do Tapirape. Mato Grosso. Brazil. 



hind RP, the transversely elevated (pinched) left 
hemitergite, the peculiar left process, the greatly re- 
duced microspiculation of the cleft membrane op- 
posite the anterior end of MF, the simple caudal 
margin of LPPT, and the peculiar form of the basal 
segment of the left cercus. 

In addition to /. amazonica, my collection 
(CAS) contains a new species from SE Brazil and 
the holotype of the older species.surinamensis 
(Ross), new combination (Pararhagadochir siirina- 
mensis Ross, 1944:433, figs. 57-60). Holotype 
male, on slide, now deposited by exchange in CAS. 
Surinam: Kwakoegron, Saramacca River, 8-VI-27 
(Cornell U. Lat 760, sub. 89). 



Ischnosembia amazonica Ross 
new species 

(Figure 12) 

Holotype. — Male, on slide, MNRJ, Data. — Bra- 
zil: Vila Amazonas near Macapa, Amapa. Matured 
in culture 17-VII-64 (E. S. Ross). 

Description. — Appearance: Small, slender, 
alate; tan with darker head and abdomen; antennae 
white-tipped, cerci entirely white. Color details (in 
alcohol): Cranium basically golden brown with 
darker pattern outlining positions of brain and 
mandibular muscles; eyes lavender black; antennal 
scape dark tan, segments 2-15 pale amber, 16 



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OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES 



lighter, distal three pure white; mouthparts varied 
shades of yellow tan. Thoracic sclerites basically 
pale yellow tan; sutures and surrounding mem- 
branes ringed with dark purple — especially on pro- 
thorax; tone of prothorax slightly darker than 
pterothorax. Legs concolorous with thorax. Wings 
pale tan with broad hyaline stripes. Abdomen basi- 
cally pale tan but appearing rust brown due to ex- 
tensive, subcutaneous mottling; tenninalia more 
yellowish, cerci entirely white. Dimensions (on 
slide): Body length 6.5 mm; forewing length 4.6 
mm, breadth 1 . 1 mm. 

Important integumental characters. — As de- 
scribed in the generic diagnosis. Of specific impor- 
tance are the size and form of the eyes, the form of 
the tergal processes, and the caudal spiculation of 
the left paraproct. 

Allotype. — Female, in alcohol with holotype 
data and disposition. 

Description: — Appearance: Small, slender; 
largely mahogany brown except for a white band be- 
tween each thoracic somite, white-tipped antennae, 
and white cerci. Color details: Cranium basically 
golden brown but heavily tinged with rust brown in 
vertex pattern. Eyes dark lavender, Antennal seg- 
ments 1-15 medium brown, segment 16 apically 
white, 17-18 white. Labrum amber yellow toward 
margins; other mouthparts varied shades of medium 
brown. All sclerotic portions of body and legs mot- 
tled mahogany brown except for transparent 
acrotergites and prescutae; all membranous surfaces 
darkly tinged with purple brown except dorsal 
membranes between thoracic somites which are 
white due to pale internal tissue. Basal segments of 
cerci whitish, tinged with dark purple toward bases; 
distal segments yellow white. Body length 9.2 mm. 

Paratypes and parallotypes. — Numerous adults 
reared from type culture, deposited in CAS, USNM, 
MCZ, MZUSP, MNRJ. and others. 

Discussion. — Ischnosembia amazonica is most 
closely related to an undescribed species from 
southern Brazil. Ischnosembia swinamensis is less 
closely related and distinguished by its more circu- 
lar head, very large eyes (eye-length greater than 
that of cranial sides) and minor differences in the 
terminalia. notably the obtusely-angulate outer flap 
of the left tergal process. I have cultured a new 
species, yet to be named, from 62 km S Ariquemes. 
Rondonia, Brazil. 




Figure 12. Ischnosembia amazonica Ross. n. sp., holo- 
type. Type locality: near Macapa, Brazil. 



Biology. — Stock for the type culture was col- 
lected in galleries spun on the undersurfaces of six- 
foot-long log sections (recently cut for charcoal 
burning). These were lying along a road cut through 
native forest near Amazon River level. The leaf lit- 
ter under the logs was very damp due to recent, 
heavy rainfall. Several other embiid species, includ- 
ing an anisembiid, were collected under the same 
logs. Adult males matured in the culture during al- 
most every month of the year 

Genus Gibocercus Szumik 

Gibocercus Szumik, 1998:141. [should have been spelled 
Gibbocerciis]. 

Type species. — Gibocercus chaco Szumik, by 
original designation. 

Distribution. — South America: Western Ama- 
zon Basin; Bolivia east of Andes, and Argentina. 

Szumik based this genus on a very limited num- 
ber of specimens. Three of her four included species 
were represented only by unique rnale holotypes 
and no associated females. I had long had this genus 
in manuscript based on more than a thousand spec- 
imens representing many species, all having males 
associated with females. 

I am dividing the genus into two subgenera, 
males of which are principally distinguished by dif- 
ferences in the basal segment of the left cercus and 



ROSS: EMBIA. BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



35 



the structure of the left tergal process (10 LP). Typ- 
ical Gibocercus has a small, echinulate nodule on 
the inner base of the left cercus (Amazonemhia 
new subgenus, does not) and a large, thin, blade- 
like inner portion of the left tergal process (in 
Amazonembia the inner portion of this process is 
narrow and thicker). 

A more detailed diagnosis of Gibocercus sensu 
stricto follows: 

Diagnosis: — Males: Large, alate, often dark 
mahogany brown, at times with prothorax and/or 
entire thorax yellow. Cranium often with a trans- 
verse, gold band between eyes (above brain); an- 
termae and cerci uniformly brown. Cranium usual- 
ly oval but at times caudally convergent; eyes small 
to large and inflated; antennae unicolorous, usually 
30-segmented; submentum quadrate, weakly scle- 
rotized, anterior margin weak, not inflexed; 
mandibles flat, thin, short with sharp, well-spaced 
apical teeth. Hind basitarsus with two ventral papil- 
lae. Tenth tergite broadly cleft to basal margin. Left 
hemitergite (10 L) large, its process (10 LP) bifid; 
inner portion long, broad, thin, blade-like, the outer 
portion shorter, narrow, acutely tapered, submem- 
branous, in some species almost obsolete. Right 
process (10 RP) an abruptly narrowed, sclerotic, 
ventrally curved talon subtended by a tiny membra- 
nous lobe. Medial flap (MF) a sclerotic "arm" pro- 
jecting forward in cleft membrane, membrane adja- 
cent to its apex not microspiculate. Epiproct (EP) 
and its sclerite well developed. Left paraproct 
(LPPT) large, well sclerotized, not fused to hypan- 
drium lobe (HP), bearing a prominent microspicu- 
late, conical lobe on its inner angle. Basal segment 
of left cercus with a very large, forward-angled, 
echinulate inner-apical lobe and a small echinulate 
nodule on its inner base. 

Females. — Not studied for generic or subgener- 
ic characters. Most are large in size and colorful, all 
have a prominent medial hind basitarsal papilla. 

Discussion. — This promises to be a very large, 
"difficuh" subgenus. To date the following species 
are named: G. chaco. G. beni and G. imicomi by 
Szumik (1998), and G. penivianus n. sp. and G. 
magniis n.sp. by me in this work. Coloration, espe- 
cially that of mature females, is very useful in 
species identification. It is regrettable that Szumik 
didn't delay publication of G. chaco until she se- 
cured a culture and reared adequate series, includ- 



ing adult females, which could easily have been se- 
cured at the type locality, near Santiago del Estero, 
only a short distance from Tucuman, her home base. 

Unfortunately, her species descriptions are too 
brief, and emphasize highly variable wing venation- 
al characters (as would be evident in a large series), 
and questionable hind basitarsal characters. 

Biology. — The many colonies I have encoun- 
tered are conspicuous, fully exposed on surfaces of 
bark, fence posts and road banks in tropical forest; 
one, G. magmis. colonized the undersurface of a 
fallen tree trunk. Occupants of colonies are highly 
gregarious. The silk is white, not obscured by cov- 
erings of pulverized debris. 

KEY TO SPECIES OF GIBOCERCUS 
SUBGENUS GIBOCERCUS 

(Males) 

1 . Body, including prothorax and abdomen, varied 
shades of brown. Pleura of abdomen almost 
concolorous with its tergites, thus not having 
conspicuous, longitudinal borders in both sexes 
(such a lack in G. chaco is assumed) 2 

— Prothorax, at times pterothorax as well, yellow 
or golden, strongly contrasting with the dark 
brown of other body sclerites. Both sexes have 
conspicuous cream white abdominal pleura ... 3 

2. As described by Szumik (1998), "Head brown 
with a dorso-posterior more less circular 
brownish area, the rest orangish [sic\ brown." 
Santiago del Estero, Argentina chaco 

— Head basically golden, longitudinally streaked 
with chestnut brown, clypeus and frons darker 
brown. Santa Cruz, Bolivia magmis 

3. Inner blade, or talon, of left tergal process (10 
LP) twice as long as broad; constricted at base. 
Mato Grosso, Brasil iiyucumi 

— Inner blade three times longer than broad; not 
constricted at base. East Andean montafia and 
yungas 4 

4. Prothorax and pterothorax bright yellow gold, 
especially dorsally. Other body sclerites brown, 
Peruvian montaiia penivianus 

— Prothorax yellow-gold, pterothorax and other 
body sclerites brown. Bolivian yungas beni 

The following three species recently described 
by Szumik (1998), apparently having very similar 
terminalia, were based on only unique holotvpes. 
Some of the characters used in the abo\e key to dis- 
tinguish them may be unreliable. 



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It is regrettable that her Bolivian species, G. 
beni, could not have been described by me at this 
time on the basis of my very large culture-reared se- 
ries of both sexes from the Santa Cruz region. As 
discussed below, under G. peniviamis n.sp., this se- 
ries can only be tentatively identified as G. beni. Fu- 
ture studies may indicate that most Gibocercus 
species compose a geographically extensive racial 
complex with coloration distinctions blending from 
locality to locality. 

Gibocercus (G.) chaco Szumik 

Gibocercus chaco Szumik. 1998:143 

Holotype. — Male, IML. Data. — Argentina: 
Santiago del Estero, Reserva Copo, 7-24-X-1990 
(J. Lopez de Cazenave) 

Coloration. — "Thorax brownish, head brown 
with a dorso-posterior more or less circular brown- 
ish area, the rest orangish [sic] brown." 

Gibocercus (G.) urucumi Szumik 

Gibocercus urucumi Szumik, 1998 p. 143. 

Holotype. — Male, MZSP. Data. — Brazil: Mato 
Grosso, Serra do Urucum-Corumba, 30-XI-1960, 
K. Lenko. 

Coloration. — "Head, 1° to 18° antennal seg- 
ment, tibiae and tarsi, and terminalia brown, the rest 
orangish [sic] brown." 

Gibocercus (C) beni Szumik 

Gibocercus beni Szumik, 1998:146. 

Holotype. — Male, MCZ. Data. — Boli\'ia: Beni 
Rurrenabaque, X-XI-1956 L. Pefia. 

Coloration. — "Antennae, prothorax, meso, 
meta-thoracic stemites yellowish; legs and termina- 
lia (except cerci) brownish, the rest dark brown; 
head with two elliptical areas; anterior one darker, 
posterior one (beUveen eyes) lighter. The abdominal 
pleurites and stemites have a longitudinal yellowish 
band." 

Gibocercus (G.) peruvianus Ross 
new species 

(Figure 13) 

Holotype. — Male, on slide, CAS. Data. — Peru: 
Yurac Plantation, 67 mi. E. of Tingo Maria (on road 
to Pucallpa) 4-X-1954 (E. S. Ross). 

Description. — Appearance: Large, alate; gener- 
ally dark mahogany brov\n with entire thorax gold- 



en yellow; appendages uniformly brown. Color de- 
tails (in alcohol): Cranium mahogany brown dorsal- 
ly with a broad transverse, yellow band between 
eyes; ventrally yellow. Eyes lavender black. Mouth- 
parts and antennae various shades of brown. Protho- 
racic and cervical sclerites clear yellow, all adjacent 
membranes cream white. Pterothorax dorsally con- 
colorous with prothorax; stemites largely yellow but 
grading to light brown caudad. All legs uniformly 
dark brown. Wings dark brown with exceptionally 
narrow hyaline stripes; cross-veins behind RP par- 
tially white. Abdomen various shades of tan except 
terminalia which are largely dark mahogany brown 
with whitish membranes; left tergal process translu- 
cent amber. Dimensions (on slide): Body length 
14.5 mm; forewing length 9.0 mm; breadth 2.2 mm. 

Important anatomical features. — As figured. 
Of specific importance is the oval cranial outline 
and rather small eyes; the exceptionally large, 
blade-like, left tergal process ( 10 LP) with a vestig- 
ial outer appendix; the extremely long, sinuous scle- 
rotic, right tergal process; the smooth, sclerotic, 
caudally-projected ventral lobe of the left paraproct; 
and the small, echinulate nodule on the inner base 
of the left cercus in addition to the large distal lobe. 

Allotype. — Female, in alcohol, with holotype 
data and disposition. 

Description. — Appearance: Large, robust; dark 
mahogany brown except for a transverse, gold 
"chevron" between eyes, whitish antennal apices 
and pale abdominal pleurae (forming a stripe on 
each side of abdomen). Color details: Cranium ba- 
sically dark reddish mahogany brown, basal pattern 
obscure; two golden areas paralleling anterior 
branches of ecdysial lines form a distinct chevron- 
like pattem between eyes. Antennal segments 1-22 
mahogany brown, 23 half white, 24—30 whitish (an- 
tennae complete). Mouthparts essentially concolor- 
ous with cranium. .Ml dorsal sclerotic portions of 
body, and legs, uniformly dark mahogany brown; 
anterior margins of thoracic scuta tan, not forming 
distinct bands; all membranous areas gray white; 
ventral sclerites paler; dorsal pleurites of abdominal 
somites 1-8 clear amber; surrounding membranous 
areas whitish, forming a pale stripe on each side of 
abdomen; terminal somites slightly darker, cerci 
uniformly brown. Body length 14.5 mm. 

Paratypes and parallotypes. — Topotypic adults 
deposited in CAS, USNM and MHNR 



ROSS; EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



37 




Figure 13. Gibocercus {Gibocercus) peruvianus Ross, n. sp., holotype. Type locality: 67 mi E of Tingo Maria, Peru. 



Discussion. — Gibocercus peruvianus appears 
to be most closely related to G. beni Szumik, per- 
haps also to G. uricumi Szumik, but, at least in the 
latter case, this cannot be determined from her short 
description. However, the more distant type locality 
of G. uricumi in Mato Grosso would appear to indi- 
cate a distinct status. Assuming that my large series 
of specimens from Bolivia's Santa Cruz region rep- 
resents G. beni, G. peruvianus is distinct in the 
bright yellow-orange of its pterothorax, that of G. 
beni is very dark brown, almost blackish. Females 
of the two species are readily distinguished by the 
overall darkness of G. peruvianus, the narrower. 



pale tan, anterior margins of the thoracic scuta 
(brighter and broader in G. beni) and a comparable 
difference in the tone of the pleural abdommal 
stripes. 

Additional records. — Peru: Colonia Perene. 18 
mi NE La Merced, Junin, 750 m estmiated, (E. S. 
Ross) CAS. Peru: Explorer's Irm, service area, Rio 
Tambopata, 30 km SW Puerto Maldonado, 290 m 
(E. S. Ross), CAS, large cultured series. 

The Perene specimens are identical to the type 
series. The more southern Tambopata series differs 
in having the pterothoracic pleurites and stemites 
medium brown instead of yellow. 



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OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES 



Biology. — Habitat is tropical rainforest along 
the eastern foothills of the Andes. The conspicuous 
colonies are most readily encountered in semi- 
cleared areas, such as coffee plantations, on sur- 
faces of logs, stumps and rotting fence posts. In cul- 
tures adults matured almost every month of the year. 

Gibocercus (G.) magmis Ross 
new species 

Holotype. — Male, on slide, CAS. Data. — Boliv- 
ia; 8 km N Santa Cruz. Matured in culture C-774, 
X-64 (E. S. Ross). 

Description: — Appearance: Very large (largest 
species of the genus); varied shades of golden 
brown throughout, prothorax concolorous with 
pterothorax. Color details (in alcohol); Cranium ba- 
sically golden, longitudinally streaked with chestnut 
brown, a pale spot present between eyes, clypeus 
and frons dark chestnut brown. Eyes lavender black. 
Antennae exceptionally long; basal segment dark 
chestnut brown, flagellar segments grading from 
medium brown to almost white at segment 33 
(complete number). Prothorax golden tan, associat- 
ed membranes dark cream. Pterothorax concolorous 
with prothorax, no pale band between the somites; 
associated membranes pinkish cream, tinged with 
rust on crests of folds. Legs varied shades of chest- 
nut brown; forelegs, especially tarsi, dark chestnut. 
Wings very large, medium brown with narrow, but 
not sharply-margined hyaline stripes; forewing's 
cross-veins: 5 RBS-RP, 4 RP-MA|+., 2 MA, +2 - 
MA3+4, and 1 MA3+4 - MP (near fork of MA); all 
cross-veins behind RP conspicuously white when 
crossing a hyaline stripe. Abdominal somites 1-8 
soft, basically cream white, heavily tinged with rust 
brown. Dimensions (on slide): Body length 17.5 
mm; forewing length 12 mm, breadth 3.0 mm. 

Important anatomical features: Cranium strong- 
ly caudally convergent; eyes large, strongly inflated, 
separated by one eye- width. Terminalia similar to G. 
pentvianus but with outer flange of 10 LP well ex- 
posed, broad-based, symmetrically and narrowly ta- 
pered, basal three-fourths sclerotic, apical fourth 
pale, its length one third that of the blade-like inner 
portion. Medial flap (MF) bulbous, sclerotic. Right 
process (10 RP) exceptionally long, sinous; sub- 
tended by a slender, transparent subprocess. Basal 
nodule of left cercus large. 



Allotype: — Female, in alcohol, CAS. Data. — A 
"sister" of the holotype. 

Description. — Very large (one of largest of all 
embiids), body length 26.0 mm. Generally chestnut 
brown throughout with pale tan intersclerotal mem- 
branes; antennae becoming pale in apical third. 
Color details; Cranium mottled chestnut brown ex- 
cept for paler muscular pattern, a pale, transverse 
zig-zag between eyes and a dark brown "cloud" in 
clypeal region. Eyes jet black. Antennal segments 
1-22 chestnut brown, 23 to 34 (complete number) 
gradually becoming pale, thence cream white to 
apex. Mandibles piceus, other mouthparts and ven- 
ter of cranium chestnut brown, submentum darker. 
Cervical and other body sclerites varied shades of 
chestnut brown; first acrotergite, meso- and 
metathoracic and abdominal sclerites somewhat 
darker; crests of all intersclerotal membranous 
folds tinged with purple; abdominal pleurites chest- 
nut brown, intervening membranes gray-pink, thus 
not forming pale, pleural, longitudinal lines as in 
some congeners. Paraprocts tan. Cerci dark purple- 
brown except for golden tips of distal segments; in- 
tersegmental membranes whitish. 

Paratypes and parallotypes. — Seven topotypic 
males and two adult females reared from culture 
774, 1964. Retained in CAS, except for the male 
deposited in IML. 

Discussion. — Females of this spectacular, large 
species are readily distinguished from G. beni, the 
other Gibocercus occurring in the Santa Cruz re- 
gion, by their larger size, uniformly brown col- 
oration, as compared to the almost black-brown fe- 
males of beni which also has contrasting tan-to- 
gold thoracic bands and pale yellow abdominal 
pleurae forming a longitudinal stripe down each 
side of the abdomen. The paraprocts are translucent 
pale cream. Cerci are very dark lavender brown 
without pale tips; the segmental joints are dark 
gray. Males are very distinct in appearance with an 
orange prothorax and a cream band between pro- 
thorax and mesothorax. The wings of G. magnus 
are especially large, light brown, with broader, ir- 
regularly-margined hyaline stripes. The wings of G. 
beni have very narrow, sharply-margined hyaline 
stripes with only three RP - MA cross-veins, (G. 
magnus has six). In the forewing G. magnus has 
two MA, +2 - MA3+4 cross-veins, (G. beni has 
none). The outer 10 LP flange of G. beni is very 
small, almost obsolete; that of G. magnus is almost 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



39 



one-third the length of the inner talon, broad-based, 
acutely tapered and desclerotized at its tip. 

Habitat. — Culture stock (C-774) was collected 
8-V-64 in second growth forest bordering an ox cart 
road in an agricultural area eight km north of Santa 
Cruz. Extensive galleries were present on the under- 
side of a fallen tree amidst weeds. Also present were 
colonies oi Pararhagadochir flavicoUis (Enderlein) 
and one or more species of Teratembiidae. Unfortu- 
nately, Cultirre 774 didn't thrive and produced only 
eight adult males and three huge adult females. 

Gibocerciis {Amazonembia) Ross 

new subgenus 

Type species. — Gibocercus {Amazonembia) 
sandrae Ross, new species 

Distribution. — South America: Upper Amazon 
drainage of Ecuador, Colombia and Peru. 

Diagnosis. — Unlike those of Gibocercus sensu 
stricto, males of this subgenus have a narrower, 
stouter, inner talon of the left tergal process (10 LP) 
and lack an echinulate nodule on the inner base of 
the basal segment of the left cercus. 

Discussion. — Like Gibocercus s. str., this prom- 
ises to be a large group with difficult-to-define an- 
atomical characters, especially in the male termina- 
lia. For this reason, such characters are figured for 
only the type species, G. sandrae n. sp. Fortunately, 
species, or at least races, can be distinguished by ap- 
parently consistent coloration of both sexes even 
when two or more species occur in the same habi- 
tats in the same locality. Besides the species treated 
below, I have also collected and reared short series 
of other species from Colombia and Ecuador which 
will not be named at this time. Because of the brev- 
ity of her description of G. (A.) nanai, I am not cer- 
tain that one of my series from the Iquitos region 
represents that species. Another series from this re- 
gion is very distinct and is given the name G. 
flavipes. If my identification of G. nanai is incor- 
rect, this would indicate that a third species of this 
subgenus occurs in the region. 

KEY TO SPECIES OF GIBOCERCUS 
SUBGENUS AMAZONEMBIA 

(Males) 

1. Distal antennal segments abruptly pure white 
3 



— Antennal segments grading from brown basal- 
ly to tan distally, i.e., not abruptly white 2 

2. Cranium dark chocolate brown with a distinct 
golden, eye-to-eye spot. Body dark brown ex- 
cept for a brilliant gold pterothorax. Upper 
Amazon of Ecuador sandrae 

— Cranium chestnut brown, eye-to-eye spot indis- 
tinct, pale cream. Body rather uniformly gold- 
en brown; pterothorax distinctly pigmented. 
Mid-Amazon region near Iquitos, Peru 
nanai 

3. Legs entirely pale yellow. Iquitos region (same 
habitat as G. nanai) flavipes 

— Coxae, trochanters and femora pale yellow, tib- 
iae and tarsi medium to dark brown. Upper 
Amazon of Ecuador (may occur in same habi- 
tat as G. sandrae) napoa 

Gibocercus {A.) sandrae Ross 
new species 

(FiGURH 14) 

Holotype. — Male, on slide CAS. Data. — Ecua- 
dor: Aliiiahui (grounds of tourist lodge), 25 km E 
Puerto Napo, Napo Prov. 475 m. Matured in culture 
30-1-95 (E. S. Ross) 

Distribution: — Central Ecuadorian montaiia, 
centered in Rio Napo tributaries. 

Name basis. — Named after my wife, Sandra, 
whose conservation efforts through land purchase, 
have insured perpetual preservation of the type lo- 
cality and its adjacent virgin forest. 

Description. — Appearance: Moderately large, 
varied shades of brown except for brilliant gold 
pterothorax. Color details (in alcohol): Cranium 
dark chocolate brown except for a broad, trans- 
verse, suffused golden band between eyes above 
brain. Eyes lavender black. Antennae basally 
chocolate brown, blending to medium brown distad. 
22-segmented (complete). Mouthparts chocolate 
brown. Cervical sclerites lemon yellow. Pronotum 
chocolate brown, faintly clouded with gold medial- 
ly; prosternum transparent light tan. Acrotergite 1, 
translucent cream white. Prothoracic and cer- 
vical membranes whitish. Pterothoracic scuta 
clear gold with contrasting brown setae; ster- 
na pale yellow. All legs chocolate brown. 
Wings brown with lavender luster, hyaline stripes 
very narrow; cross-veins: 2 RBS - RP. 2 RP - MA,^;- ■''o 



MA, 



MA 



■3+4' 



1 MA - MP; cross-veins behind 



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OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES 




TERMINALIA 

VENTRAL 
Figure 14. Gibocercus {Amazonembia) sandrae Ross, n. sp., holotype. Type locality: 25 km E of Puerto Napo, Ecuador. 



RP bordered with white when crossing a hyaline 
stripe. Abdomen, except terminalia, whitish but 
strongly tinged with blackish brown, anterior mar- 
gins of tergites narrowly dark brown; ninth tergite 
dark chestnut brown medially, becoming piceous 
laterally; tenth tergite, its processes, epiproct and 
sclerotized portions of cerci glossy dark brown, 
piceous at margins; membranous areas cream 
white. Dimensions (on slide): Body length 15.0 
mm; forewing length 8.5 mm, breadth 1.6 mm. 

Important integumental features. — As figured. 



Allotype. — Female, in alcohol (CAS). Data. — 
From holotype 's culture. 

Description. — Large, body length 17.0 mm. 
Black except for white bands between thoracic 
somites, and a longitudinal pleural stripe on each 
side of abdomen, and a golden area across cranium. 
Color details (in alcohol): Cranium blackish brown 
except for large, marginally suffused, orange band 
across head between eyes (above brain) and anten- 
nal bases. Antennal segments 1-18 dark brown, 19 
becoming white apically, 20-24 pure white (com- 
plete). Dorsal body sclerites jet black except for 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



41 



narrow white anterior margins of meso- and metas- 
cuta of thorax; combined with white membranous 
areas, two conspicuous white thoracic bands are 
formed. Legs largely dark chocolate brown, or 
black; femur tibial joints golden. Abdominal ter- 
gites glossy black, margins of tergites 1-7 golden; 
pleural membranes white, thus forming a longitudi- 
nal white line on each side of abdomen; stemites 
smokey light brown, paragenitals darker. Para- 
procts pale, clouded with light brown. Basal seg- 
ments of cerci lavender black, distals medium 
brown; joint membranes lavender. 

Paratypes and parallotypes. — Hundreds of cul- 
tured topotypic males and females. Deposited in 
CAS, USNM, MCZ, NMQ, MZUSP and MNHP 

Biology. — Large colonies are very abundant on 
most trees on the grounds of Cabaiias Aliiiahui and 
less so in the adjacent primary forest. They are very 
conspicuous because the silk isn't obscured by pul- 
verized habitat material. The congener, G. {A.) 
napoa n. sp., distinguished by its white-tipped an- 
tennae and other features, occurs in virgin forest, 
along the road between Puerto Napo and Puyo, and 
elsewhere in the region. 

Gibocercus {A.) napoa Ross 
new species 

Holotype. — Male, on slide, CAS. Data. — Ecua- 
dor: 3 mi N Puyo, Napo-Pastaza, 950 m, III-55 (E. 
S. Ross). 

Name basis. — Relates to type region. 

Description. — Appearance: Rather slender, 
alate, medium sized (body length 12.0 mm); entire 
thorax and legs largely pale yellow, head, wings and 
abdomen dark brown. Color details (in alcohol): 
Cranium dark chocolate brown with a small, faint, 
pale maculation between eyes; narrow pale rim 
around black eyes; ventrally becoming medium 
brown. Antennal segments 1-15 blending from 
dark brown to tan, 17 to 26 abruptly pure white 
(complete number). Thorax entirely pale yellow 
with dark brown setae, prothorax slightly darker 
(more golden); membranes and axillary area of 
wings cream white, almost concolorous with scle- 
rites. Wing bands brown; hyaline stripes very nar- 
row, sharply margined; in forewing 4 RP - MA, +2, 
no MA, +2 - MA3+4, 1 MA - MP cross-veins, all 
conspicuously white. Coxae, trochanters and femo- 
ra of all legs concolorous with thorax; all tibiae and 



tarsi medium to dark brown. Abdomen mottled 
dark brown; tenth tergite, its processes, and left cer- 
cus much darker; other cercus segments medium- 
brown, including joints. Dimensions (on slide): 
Body length 12.0 mm; forewing length 7.5 mm, 
breadth 2.0 mm. 

Important anatomical features: Cranium and 
mandibles similar to G. {A.) sandrae, outer margins 
of mandibles evenly arcuated. Terminalia similar to 
G. (A.) sandrae but with base of 10 LP broader. 

Allotype. — Female, in alcohol, CAS. Data. — 
From holotype culture. 

Description. — Appearance: Very dark brown 
with pale intersegmental membranes, an orange 
area between eyes, antennae white-tipped. Color 
details (in alcohol): Cranium black blending to dark 
orange in a large, transverse "cloud" between eyes 
(above brain). Mouthparts and antermal, segments 
1-21 brown, others (22-27) pure white. All tho- 
racic sclerites almost black; all membranes gray 
white, forming a narrow pale band between each 
somite. All leg segments dark chocolate brown. Ab- 
dominal sclerites likewise almost black, including 
the caudal tergites; pleural membranes grayish, 
forming a pale longitudinal band on each side. 
Basal segments of cerci medium brown, apicals tan. 
Body length 14.5 mm. 

Paratypes. — 18 topotypic males, CAS and 
NMQ. 

Additional records (CAS): 1 male, Ecuador: 
Sacha Pacha, Rio Aguarico, 200 m, matured XII-91 
(E. S. Ross); numerous juvenile, females and 
males, 2-5 km N. Puyo, Napo-Pastaza, 950 m, II- 
55 (E. S. Ross) colonies on stumps in pasture. 

Gibocercus {A.) nanai Szumik 

Gibocercus nanai Szumik, 1998:147. 

Holotype. — Male, USNM. Data. — "Peru, Lore- 
to, Callicebus Res. Station, Mishana, Rio Nanay, 25 
km SW Iquitos, 10-17-1-80, S.B. Heppner." 

Discussion. — Unfortunately this species was 
incompletely described from a single specimen. 
When I visited the Iquitos region in 1 983 I secured 
cultures of two very distinct new species of Gibo- 
cercus which I have long had in manuscript pend- 
ing an occasion to comprehensively treat them 
within a new genus. Szumik's color description. 
"General coloration brownish, 1 6° antennite to the 



42 



OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES 



tip yellowish brown" is barely sufficient to distin- 
guish G. (A.) nana! from the other Iquitos species, 
G. {A.) flavipes n. sp., which has pure white anten- 
nal apices, and the entire thorax and all legs pale 
yellow, as well as other distinct features. If Szu- 
mik's specimen had such distinctive coloration, she 
surely would have noted it. It is significant that the 
two species occur in the same habitat. I will now 
offer a more detailed color description of what I as- 
sume to be G. {A.) nanai based on my extensive se- 
ries (CAS). 

Plesiotype. — Male, on slide, CAS. Data. — 
Peru: Amazon Camp, Rio Momon, near Iquitos, 
matured in culture 11-83 (E. S. Ross). 

Description. — Appearance: Medium sized, gen- 
erally yellow brown, head chestnut brown, eyes 
contrastingly black, antennae grading from chest- 
nut brown at base to cream at apex; all legs cream. 
Color details (in alcohol): Cranium evenly chestnut 
brown, broadly paler from eye-to-eye (above brain). 
Eyes jet black. Antennae grading basally from 
chestnut brown to cream at apex. Mouthparts light 
chestnut brown. Body sclerites varied shades of 
chestnut brown on a yellowish base; all intervening 
membranes white, forming a conspicuous pale 
band between pro-and mesothorax. All coxae, tro- 
chanters, and femora cream white, tibia and tarsi 
grading to tan; foretarsi darker. Wings light brown, 
hyaline stripes not sharply defined. In forew^ing. 
one cross-vein between RP - MA 1^2- none between 
MA, +2 and MAj^j, one MA - MP (midway on 
MA); cross-veins indistinctly white. Sclerotic por- 
tions of terminalia golden with amber margins; 
inner talon of left process (10 LP), and margins of 
more brownish basal segment of left cercus, dark 
amber; basal segment of left cercus pale tan, distal 
segments grading distad from pale tan to cream 
white. Dimensions (on slide): Body length 9.0 mm; 
forewing length 6.2 mm, breadth 1.5 mm. 

Important integumental characters. — Cranium 
caudally convergent, sides about one-eye-length 
long. Eyes large, inflated; interspace one and one- 
half eye-widths. Medial flap (MF) of terminalia 
globose, smoothly sclerotized. Talon of right tergal 
process (10 RP) slender, elongate, darkly sclerotic. 
Lobe of left paraproct (LPPT) conical. Left cercus 
lobe especially long, projected diagonally forward; 
apical cercus segments exceptionally long, slender. 

Szumik used as a key character (1998:142) the 



absence of a "membranous band" between the men- 
tum and submentum. Slide preparations reveal that 
these structures are clearly separated by a membra- 
nous interval in all species of the order. 

Female, in alcohol, CAS. Data. — Same as ple- 
siotype. 

Description. — Appearance: Cranium, prothorax 
fore and mid legs gold to light amber, remainder of 
thorax and most of abdomen shades of mahogany 
brown to medium brown; abdominal segments 8-10 
pale yellow, cerci light tan. Color details: Cranium 
orange, paler between eyes; anterior tentorial pits 
piceous; venter of head and mouthparts orange. 
Eyes jet black. Basal antennal segment amber, seg- 
ments 2-20 dark brown. 21-28 pure white. Prono- 
tum dark amber, edges darker; acrotergitel clear 
light amber. Mesonotum basically dark amber, 
heavily clouded with dark mahogany brown, anteri- 
or margin cream. Metanotum similar but more 
evenly mahogany, anterior cream margin narrower. 
Acrotergite 2 amber. Forelegs entirely light amber; 
midlegs similar but coxae are mahogany brown; 
hind legs with coxae, trochanters and basal two- 
thirds of femora mahogany brown, blending distad 
to pale amber; tibiae and tarsi light amber Abdom- 
inal tergites 1 and 2 lighter mahogany brown, others 
caudad becoming lighter and cream white at sides, 
combining with pale pleurae to form pale lateral, 
longitudinal stripes; somites 8-10 abruptly nar- 
rowed and pale yellow. Venter of body anteriorly 
pale yellow becoming tan caudad; paragenital ster- 
nites tan at sides. Paraprocts translucent cream 
white. Basal segments of cerci light tan, distal seg- 
ments paler. Body length 13.5 mm. 

Available specimens. — Twenty-six adult males 
and three adult females with plesiotype data depos- 
ited in CAS. USNM, and MHNR 

Gibocercus {A.) flavipes Ross 
new species 

Holotype. — Male, on slide, CAS. Data: Peru: 
Amazon Camp, Rio Momon, near Iquitos, 6-XII-82 
(E. S. Ross). 

Name basis. — Refers to entirely yellow legs of 
males. 

Description. — Appearance: Medium sized 
(body length 13.0 mm); dark mahogany brown ex- 
cept for white tipped antennae, thorax and legs en- 
tirely pale yellow, except for brown tarsi. Color de- 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



43 



tails (in alcohol): Cranium mahogany brown except 
for diffuse, broad "cloud" between eyes (above 
brain). Eyes lavender black. Antennal segments 
1-16 mahogany, 17 yellow brown with white apex, 
18-24 pure white including setae (complete num- 
ber). Mouthparts light mahogany, ventral area of 
head golden brown. Cervical and thoracic sclerites 
pale yellow, pronotum slightly darker; all associat- 
ed membranes cream white. All legs entirely pale 
yellow except for brown tarsi. Wings medium 
browTi except for white axillary region, narrow hya- 
line stripes. Forewings with one short cross-vein 
(not white) between RBS and RP, two white RP - 
MAi^2' one between MA ,+2 and MA3+4, and one 
white one between fork of MA and MP. Abdominal 
somites 1-9 gray white heavily mottled subcutane- 
ously with rust brown: somite 10 and base of left 
cercus glossy mahogany brown blending to 
piceous, apices of 10 RP and LC,, lobe and medial 
sclerite golden brown; membranes pink-white; 
other cercus segments yellow tan. Dmiensions (on 
slide): Body length 13.0 mm; forewing length 8.5 
mm, breadth 2.0 mm. 

Important anatomical characters. — Cranium 
elongate-oval, sides behind eyes almost four eye- 
lengths long, broadly arcuate, scarcely convergent. 
Eyes small, strongly inflated; interspace about four 
eye-widths wide. Medial flap (MF) rather flat, mi- 
crostrigose. Talon of right tergal process (10 RP) 
short. Left paraproct (LPPT) lobe broadly rounded. 
Left cercus lobe relatively short, apically narrowed, 
almost conical. 

Allotype. — Female, in alcohol, CAS. Data. — 
From holotype's culture. 

Description. — Appearance: Various shades of 
mahogany brown, pronotum darkest; membranous 
areas cream white. Cranium with a bright yellow 
interocular spot: antennae white-tipped. Color de- 
tails (in alcohol): Cranium basically mahogany 
brown except for a very conspicuous, broad, dif- 
fused yellow band between eyes (above brain) and 
darker pigment pattern associated with internal 
muscle attachments. Antennal segments 1-19 ma- 
hogany brown, 20-28 pure white. Mouthparts and 
ventral area chestnut brown, clypeolabral mem- 
brane cream. Pronotum piceous brown, other tho- 
racic tergites, including acrotergite 1, varied shades 
of dark mahogany brown; intersclerotal membranes 
cream, thus forming two intersomital pale bands 
on thorax. All legs varied shades of chestnut 



brown, fore and mid tarsi tan, hind tarsi cream. Ab- 
dominal tergites mottled mahogany brown with a 
conspicuous cream stripe on each side. Ventral body 
sclerites shades of translucent tan, except for chest- 
nut brown presternum; paragenital sternites black- 
ish brown, except medially cream on segment 8. 
Paraprocts translucent pale tan. Basal segments of 
cerci dark tan, distals pale tan. Body length: 16.0 
mm. 

Paratypes and parallotypes. — Eleven males and 
three topotypic females, CAS, USNM, MHNR 

Genus Pararhagadochir Davis 

Pai-arhagadochir Davis, 1940a: 181; 1942:1 14.— Ross, 
1944:420; 1972:133; 1992:123.— Szumik, 1996:51; 
1998:141 (cladogram). 

Type species. Embia trinitatis Saussure, 1896, 
by original designation. 

Distribution. South America (at least one spe- 
cies, P. trinitatis, has spread, probably by means of 
commerce, into Central America). 

Diagnosis. Males: Small to moderately large (6- 
14 mm), slender; usually richly pigmented and mel- 
anized. Head and eyes variable in form; antennae al- 
ways unicolorous (never white-tipped); mandibles 
oligotomoid, apices dentate; submentum almost al- 
ways sclerotic, shield-like, sides and anterior mar- 
gin usually infle.xed. Almost always alate (apterous 
or short-winged males occasionally occur in nor- 
mal-winged species) vannal area reduced; usually 
no cross-veins in wings behind MA. Hind basitarsus 
with second papilla greatly reduced, often only a 
tiny membranous flat spot not visible from lateral 
aspect, apparently absent in males of some species. 
Tenth tergite broadly cleft to base. Left process (10 
LP) almost always bifid, composed of an inner talon 
and a flat, often submembranous, outer portion 
(flange). Right process (10 RP) composed of a ven- 
trally-directed talon, subtended by a small, sub- 
membranous, ventral nodule. Medial flap (MF) well 
developed, often a striated sclerotic arm extending 
almost to base of cleft; forward end often flared and 
directed toward, or into, a microspiculate pouch in 
the cleft membrane. Epiproct (EP) very large, cau- 
dally acute; its sclerite long and conspicuous. Left 
paraproct (LPPT) large, its caudal margin of two 
types: a trinitatis-type with the inner angle pro- 
duced as a small spine and a teniiis-typs with a me- 
dial, microspiculate, rounded nodule and no inner 



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OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES 



spine. Basal segment of left cercus with a single, 
large, globose, densely-echinulate, inner-apical 
lobe; distal segments of cerci never white. 

Females: With coloration similar to males. Ge- 
neric anatomical characters not investigated. Sec- 
ond hind basitarsal papilla well developed. 

Discussion: Except for two large new species 
from SE Brazil, and a very small new species from 
NE Brazil, which have a soft submentum, males of 
Pararhagadochir may be recognized by their scle- 
rotic shield-like submentum in combination with an 
almost always bifid left process and a microspicu- 
late membranous pouch opposite the forward end 
of a rod-like medial flap of the terminalia. The new 
genus Litosembia has these characters but differs in 
its very small size, minute 10 LP talon, and many 
other characters. 

As suggested by at least thirty-five mostly new 
species in my collection (CAS), Pararhagadochir 
is the largest genus of New World Embiidae. At this 
time I will only review the named species and de- 
scribe two interesting new ones. For convenience, 
figures of known species scattered in the literature 
are republished. Fortunately, I have been able to 
study the types of most of these. The genus prom- 
ises to be "difficulf " and must be studied on the 
basis of large, cultured series, with special consid- 
eration given to the coloration of females. 

The various species occur in a wide range of 
habitats: on bark, road banks and under stones; 
from deserts to high paramo and cloud forests up to 
more than 3000 meters altitude. 

KEY TO SPECIES OF PARARHAGADOCHIR 

(Males) 

1. Inner comer of left paraproct (LPPT) with a 
minute, sharp or blunt spine; caudal margin 
without a microspiculate nodule 2 

— Inner comer of LPPT without a spine, caudal 
margin often with a microspiculate nodule ... 6 

2. Outer flange of left tergal process (10 LP) 
long, acutely tapered, often apically twisted. N 
South .America trinitatis complex 

— Outer flange of 1 LP short, apically rounded, 
or irregular in shape 3 

3. Base of 10 LP greatly elongated, narrow; at 
least three-fourths of the process' length. Ar- 
gentina trachelia 



— Base of 10 LP shorter, not exceeding half the 
length of the process. Widespread species 4 

4. Exceptionally large, long-winged species, 
spine of left paraproct blunt. Machu Picchu, 
Peru picchua 

— Average or small sized species, spine of para- 
proct sharp. Lowland habitats 5 

5. Apterous. Body and legs pale, translucent 
cream white; head and antennae chestnut 
brown. Northern Argentina pallida 

— Winged. Body and legs uniformly brown; pro- 
thorax often yellowish, or golden. Widespread 
species flavicollis complex 

6. Medial flap (MF) of tenth tergite dorsally scle- 
rotized. strongly arched; left process (10 LP) 
with outer flange very short, narrow; talon of 
right process (10 RP) minute. Lower Amazon, 
NE Brazil christae 

— Medial flap narrow, sclerotized only on ventral 
surface, not arched; flange of 10 LP broad, al- 
most as long as inner talon; talon of 10 RP 
large, conspicuous 7 

7. Caudal margin of left paraparoct (LPPT) 
smooth, without a microspiculate nodule 1 1 

— Caudal margin of LPPT with a microspiculate 
nodule 8 

8. Talon of left tergal process (10 LP) evenly 
curved outward. Amazon Basin .... bicingillata 

— Talon abruptly turned leftward, almost at 90°. 
Parana Basin and southward 9 

9. Head broadly circular in form. Eastem Para- 
guay, Argentina and S Brazil confusa 

— Head elongate-oval, sides behind eyes short, at 
times strongly convergent 10 

10. Eyes very large, separated by only one eye- 
width; sides behind eyes short, less than length 
of an eye. strongly convergent. Santa Cruz re- 
gion, Bolivia tenuis 

— Eyes small, separated by about four eye widths; 
sides behind eyes long, at least two eye-lengths 
long, only slightly convergent. Bolivia and Ar- 
gentina birabeni complex 

1 1 . Submentum strongly sclerotized, all margins 
inflexed. SE Brazil balteata 

— Submentum weakly sclerotized, anterior mar- 
gin not inflexed weak. NE Brazil minuta 



ROSS: EMBIA. BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



45 



Pararhagadochir trinitatis (Saussure) 

(Figure 15) 

Embia trinitatis Saussure, 1896a:293; 1896b:352, fig. 
13.— Enderlein, 1912:52, \06 (trinitatensisisiclM).— 
Navas, 1918:99. 

Oligotoma trinitatis (Saussure), Krauss, 191 1:42. 

Pararhagadochir trinitatis (Saussure), Davis, 1940a: 182; 
1942:114.— Ross, 1944:421; 1992:124.— Szumik, 
1998:141 (cladogram). 

Oligotoma flavicollis Krauss. 1911:43 (not Embia flavi- 
collis Enderlein). — Enderlein, 1912:100 (as a syn of 
flavicollis Enderlein, in error) . — Ross, 1944:422 (as 
syn. oi trinitatis Saussure). 

Type. — A cotype redescribed and figured by 



Davis (1940a), deposited in Mus. d'Hist. Nat, 
Geneva, Switzerland. Data. — Trinidad probably 
Port of Spain (Urich). 

Discussion. — Based on hundreds of reared 
specimens (CAS) from Trinidad, Venezuela, Co- 
lombia, Panama, Costa Rica, Guatemala and Hon- 
duras, this appears to be a very widespread, variable 
species. At least in Central America, its range has 
been extended in human commerce. In many local- 
ities males are uniformly brown. One series from 
relatively high altitude in Venezuela, near Santo 
Domingo, Barinas, 1350 m, appears to represent 
a distinct race or species. For the present, how- 




TERMINALIA 

VENTRAL 

Figure 15. Pararhagadochir trinitatis (Saussure). Type locality: Trinidad. Specimen fi-om near Azulita. Merida. 750 
m, Venezuela (may be a subspecies). 



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ever, it is best to treat males of all northern South 
American Pamrhagadochiv having a tapered outer 
flange of 10 LP as P. trinitatis. 

Records (all CAS, collected by me unless other- 
wise indicated). — Trinidad: numerous localities. 
Venezuela: Rancho Grande (granite road bank); 10 
km S of Rancho Grande; 25 km SW Socopa, Bari- 
nas, 200 m; 5 km NW Azulita, Merida, 750 m; 6 
km N Michelina, Tachira, 1200 m; near Santa Mar- 
ia, N of Caripe, 250 m; 36 km N Bocono, Trujillo. 
2225 m; Guanaguana, Monaga, 350 m; 3 km SE 
1350 m and at 1550 m, 8 km SE of Santo Domin- 
go, Barinas (both in paramo, may represent a dis- 
tinct, high altitude species). Colombia: 4 mi SE 
Villavicencio. Meta, 410 m; San Sebastian de 
Rabaga, Sierra Nevada Santa Marta, 2000 m 
(Borys Malkin). Panama: Cerro Campana. COSTA 
RICA: Finca la Lola, near Sequirres; 5 mi S Peiia 
Blanca. Honduras: 23 mi. NE Talanga, 1700 ft 
(O'Brien). Guatemala: San Jose (coastal port). 

Biology. — This species fonns conspicuous col- 
onies, especially on surfaces of road and trail 
banks, in diverse habitats from sea level tropical 
forest to paramo as high as 2225 m, and in deserts. 
Obviously a complex of difficult-to-define species 
and races awaits intensive study by a qualified res- 
ident specialist. 

Discussion. — My notes on the synonym Oligo- 
toma flavicollis Krauss, 1911, are as follows: 
Type. — Male, in fragmentary condition, deposited 
in the Zoologischen Museum der Univ. Berlin. La- 
bels. — "Orinoco, Moritz"; "2737"; "type" (red); 
"Olyntha flavicollis Krauss Male Type!"; "Rha- 
gadochir flavicollis Endl. Male Dr Enderlein det 
1911." 

The above type consists only of the pterothorax, 
two wings, and the first three abdominal somites. 
Krauss' description and figures covers more ana- 
tomical details and it is therefore evident that por- 
tions of his specimen were subsequently lost. A 
right forewing glued by a curator to the mesothorax 
belongs to some other species, apparently Embia 
fibtilatoria Enderlein of which the Berlin Museum 
has a series. The right wing, however, attached to 
the type of P. Jlaricollis clearly indicates that the 
specimen is a Pararhagadochir, one with venation 
identical to that of P. trinitatis. 

Even if an Orinoco population (somewhere on 
that long river) proves to be racially distinct, the 



name P. flavicollis Krauss, 1911, will not be avail- 
able for it is a secondary homonym of Pararha- 
gadochir flavicollis (Enderlein), 1909, of Bolivia. 
My examination of Enderlein 's type indicates that it 
represents a distinct species and it is not a sub- 
species of P. trinitatis as suggested in my 1944 re- 
vision. 

Pararhagadochir flavicollis (Enderlein) 

(Figure 16) 

Embia flavicollis Enderlein. 1909:184.— Krauss, 1911: 
68.— Navas, 1918 p. 100. 

Rhagadochir flavicollis (Enderlein), Enderlein. 1912:56, 
100. 

Pararhagadochir flavicollis (Enderlein). Davis. 1940a: 
183. 

Pararhagadochir trinitatis flavicollis (Enderlein). Ross, 
1944:424 ( in error as a subsp. of trinitatis). 

Pararhagadochir schadei Ross. 1944:427. Holotype: 
Male, on slide. MCZ; data: Villa Rica, Paraguay, De- 
cember (E Schade). New synonym. 

Lectotype (by present designation). — Male, on 
slide, deposited in the Polska Akademia Nauk, War- 
saw. Labels. — "Bolivien Prov. Sara" [Dept. Santa 
Cruz, about 100 km NW city of Santa Cruz], 
"Embia flavicollis Ended. Type det. Dr. Enderlein." 

Description (of lectotype). — Appearance: Alate, 
rather small, slender; prothorax exceptionally nar- 
row, cream yellow in contrast to dark brown head, 
body, and wings; wings unusually large in propor- 
tion to body size. Color details (dry): Cranium dark 
chocolate brown, edges of clypeus and anterior apo- 
physeal pits, as well as venter, dark amber. Eyes 
piceous; antennae medium brown throughout, ex- 
treme apical segments lost. Mandibles pale amber 
with reddish amber incisor margins; submentum 
golden brown. Pronotum, its pleura, membranes 
and cervical sclerites cream yellow; prostemum 
golden brown. Pterothorax and abdomen mahogany 
brown; all leg segments similar but with coxae and 
trochanters yellow brovra, forecoxae darker. Ab- 
dominal terminalia, including cerci, mahogany 
brown. Dimensions (on slide): Body length 8.5 
mm.; forewing length 6.5 mm, breadth 1 .6 mm. Im- 
portant anatomical characters as figured. 

Discussion. — My knowledge of P. flavicollis is 
based on a study of its lectotype. here figured, and 
large, almost topotypic series I reared from the 
Santa Cruz region of Bolivia. The lectotype of P. 
flavicollis closely resembles the holotype of P. 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



47 




Figure 16. Pararagadochir flavicoUis (Enderlein). Lectotype. Type locality: Santa Cruz region. Bolivia. 



schadei and this justifies the latter "s synonymy. 

Records (all CAS). — Bolivia: Santa Cruz and 
Angostura (70 km SW Santa Cruz). Argentina: 
Corrientes; Tres Isletas, Chaco. Paraguay: Villa 
Rica (Schade). 

Pararhagadochiv flavicoUis is the first named 
of a large, diverse complex of species related to P. 
trinitatiis but usually smaller and always having the 
flange of 10 LP short and distally rounded instead 
of long and tapered. Many of the species and/or 
races occur in the Andes and Amazon and Parana 
basins. Only two new species representing ex- 
tremes in the diversity of the P. flavicoUis complex 
are described at this time. 

Pararhagadochir pallida Ross 
new species 

(Figure 17) 

Holotype. — Male, on slide. IML. Data. — Ar- 
gentina: Salta, Cabeza de Buey, matured in culture 



(C-756), 30-XII-64 (E. S. Ross). 

Description. — Appearance: Apterous, very 
small (body length 8.5 mm), body and legs entirely 
translucent creain white, head and antennae con- 
trastingly chestnut brown. Color details (in alcohol): 
Cranium uniformly chestnut brown, without pat- 
tern. Eyes black. Basal two antennal segments dark 
brown, all others uniformly medium brown. 18 seg- 
ments. Mandibles pale amber, dentation dark am- 
ber; other mouthparts. including submentum and 
ventral bridge, translucent yellow tan, palpi medium 
brown. Body uniformly translucent cream white, 
legs slightly darker; terminalia concolorous with 
body except for amber left tergal process and pale 
brown cerci (except for pale basal segment of right 
cercus). 

Anatomical characters. — As figured. Medial 
flap (MF) resembles a concave disc on edge; talon 
and flange of left tergal process (10 LP) almost 
equal in length; talon of right process minute, adja- 



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OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES 




TERMiNALIA 
VENTRAL 



Figure 17. ParwhagadochU- pallida Ross, n. sp.. holot>pe. Type locality: Cabeza de Buey. Salta, Argentina. 



cent ventral nodule larger than talon; left paraproct 
caudally de-sclerotized. inner corner bearing a 
sharp, acute projection. 

Allotype. — Female, in alcohol, IML. Data, — 
reared in holotype's culture. 

Description. — Appearance: Pale, only slightly 
darker than male. Color details (in alcohol): Crani- 
um basically chestnut brown with pattern outlined 
in rust brown. Eyes black. Antennae cream white. 
Pronotum clear pale yellow, all other body sclerites 
pale amber but darkened by tinges of rust. Legs 
similar but with femora more heavily tinged. Cerci 
with basal segments tinged with rust, apicals paler. 
Body length 10.0 mm. 

Paratypes and parallotypes. — Numerous adults 
of both sexes deposited in CAS, USNM, BMNH 
and MZUSP 

Discussion. — This remarkable pale species 
with terminalia very^ similar to P. Jlavicollis and P. 
trachelia, exhibits no color variation in my large 



cultured series. It was collected in a broad valley 
with low thorn forest at the site of a road mainte- 
nance camp. Embiid colonies were common under 
small stones and bricks in the thin shade of low, 
thorny trees. Also at this site: Pararhagadochir tra- 
chelia (Navas), P. birabeni (Navas) and Chelicerca 
tigre Szumik. 

Pararhagadochir picchua Ross 
new species 

(Figure 18) 

Holotype. — Male, on slide, CAS. Data. — Peru: 

Machu Picchu. Matured in culture 23-XII-82 (E. S. 
Ross). 

Description. — Appearance: Very large (body 
length 14.0 mm), wings disproportionally large; 
body varied shades of chestnut brown with piceous 
sutural lines, head much darker than body. Color 
details (in alcohol): Cranium dark mahogany 
brown with darker pattern outlining muscle attach- 
ment areas and the clypeus. Eyes lavender black. 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



49 




Figure 18. Pararhagadochir picchua Ross, n. sp., holotype. Type locality: Machu Picchu, Peru. 



Basal antennal segment mahogany brown, all oth- 
ers to apex (segment 25) medium brown with 
cream joints. Venter of cranium and mouthparts 
golden brown, palpi darker. All thoracic sclerites 
and legs light chestnut brown with piceous sutural 
lines and edges; membranous areas cream white. 
Abdomen similar with cream white pleura forming 
pale lateral bands; terminalia darker, distal cercus 
segments pale tan. Dimensions (on slide): Body 
length 14.0 mm; forewing length 12.0 mm, breadth 
2.5 mm. 

Anatomical distinctions. — Cranium elongate- 
oval, sides behind eyes convergent; eyes bulging, 
separated by two and one half eye-widths; mandi- 
bles elongate; submentum with arcuate sides, apical 
comers produced forward. Forewings with RP - 
MA fork near wing base, 5 RP - MA ,+2 cross- 
veins, 2 RP - MA2+3 cross-veins, no others behind 
these. Hind basitarsus with very dense setae, medi- 
al papilla absent. Terminalia as figured, with base 
of left tergal process (10 LP) exceptionally broad 
and short; inner talon feebly arcuated, narrow, very 



sharp; flange almost equal in length, narrow at 
base, then expanded distad, apex asymmetrically 
rounded. Spine at inner corner of left paraproct 
sclerotic but apically round not sharp as in related 
species. Talon of right process short, small, direct- 
ed ventrad, not visible from above. 

Allotype. — Female, in alcohol, CAS. Data. — 
From holotype 's culture. 

Description. — Appearance: Large (body length 
18.0 mm), blackish brown with whitish intersclero- 
tal membranes; femoro-tibial joints cream white. 
Color details: Cranium entirely blackish brown ex- 
cept for very faint pattern. Eyes black. Basal anten- 
nal segment concolorous with cranium, all other 
segments to apex (segment 26) uniformly chocolate 
brown. All mouthparts dark mahogany browni. All 
body sclerites blackish brown except acrotergites 
which are dark chestnut, abdominal pleurites 
amber; all intersclerotal membranes whitish, form- 
ing a pale longitudinal pleural stripe on each side of 
abdomen. Legs and venter of body varied shades of 



50 



OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES 



brown; hind trochanters pale amber, venter of hind 
femora shades of brown to tan. Basal segments of 
cerci almost black; apical segments extremely long, 
chestnut, blending distad to cream at extreme tip. 

Paratypes and parallotypes. — Numerous cul- 
tured males and a few adult females deposited in 
CAS, MHNP, USNM and BMNH. 

Discussion. — Males of this species, like those 
of so many other high altitude embiids, have very 
large wings in proportion to a smaller slender body. 
There are also distinctions in the tenninalia. Farther 
north in the Andes I collected several related new 
species, one at over 3000 m elevation in paramo. 

At Machu Picchu I encountered small colonies 
of this species in dense moss growing on trail 
banks. They were extremely rare, difficult to locate. 

Pararhagadochir trachelia (Navas) 

(Figure 19) 

Rhagadochir trachelia Navas, 1915:135. 

Embia ti-achelia (Navas), Navas, 1918:100; 1922:363 
(record); 1923:197 (record); 1924:10 (records); 
1930:72 (record). 




Figure 19. Pararhagadochir trachelia (Na\'as). Type lo- 
cality: Prov. Santiago del Estero, Argentina. 



Pararhagadochir trachelia (Navas), Davis, 1940a: 184, 
figs. 51-66 (ex parte). — Ross, 1944:424 (redesc. 
type).— Szumik, 1998:141 (in cladogram); 1999:81 
(biology). 

Lectotype (by present designation, but speci- 
men not so labeled). — Male on slide. La Plata Mu- 
seum, Argentina. Labels. — "Rep. Argentina, Pr. 
Santiago d. Estero 190- C. Bruch." Another 
"Typus" with the above data is in the Navas collec- 
tion (Barcelona). 

Discussion. — Pararhagadochir trachelia, ap- 
parently restricted to Argentina, is readily recog- 
nized by the very narrow, greatly elongated base of 
10 LP terminated by a short, apically rounded 
flange and a twice-as-long, evenly-tapered talon. 
Also, the inner spine of LPPT is exceptionally 
small. In some populations the prothorax is yellow- 
ish, in others it is brown (concolorous with the 
pterothorax). There is also much variation in body 
size from locality to locality and wings may be 
present or absent. However, the latter condition 
should never be the basis for proposing new species 
or racial names. 

The following is a list of populations I have 
sampled during two trips to Argentina (1951 and 
1964): Salta: Cabeza de Buey, males alate. Jujuy: 5 
mi N Jujuy, males apterous, under stones; 10 km S 
Yuto, small, apterous and alate males in the popula- 
tion. Catamarca: 20 mi W Lavalle, large apterous, 
bicolorous males, colonies in matted leaves on a 
ridge crest; Tucuman: NW of Villa Padre Monti 
(Km 53 N of Tucuman), alate males bicolorous; 
Mendoza: 10 km E Villavicencio, under stones in 
extremely dry desert; males alate, unicolorous 
brown, base of 10 LP exceptionally long. 

Pararhagadochir bicingillata 

(Enderlein) 

(Figure 20) 

Oligotuma bicingillata Enderlein, 1909:191; Krauss, 
1911:45.— Enderlein, 1912:52, 93.— Navas, 1918: 
102.— Davis, 1940c:384 .—Ross, 1944:497. 

Pararhagadochir bicingillata (Enderlein), Ross, 
1972:138 (new combination). 

Pararhagadochir davisi Ross, 1944:432. figs. 52-56 
(Type male, MCZ. Data: "Brasil: Parintins, Oct. 2 
(Parish)"; Ross, 1972:138. New synonym. 

Type. — Mature female, originally pinned, now 
on microscope slide (my preparation). Deposited in 
the Polska Akademia Nauk, Warsaw. Labels. — 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



51 



"Para" [Brazil]. "Type" [red], "Oligotoma 
bicingillata Enderl. Female Type det. Dr. Ender- 
lein." 

Discussion. — Prior to Ross. 1972. all references 
to this name simply cited Enderlein "s type. 1 have 
studied this specimen and noted that it is a badly 
damaged female of an endemic species represent- 
ing a valid name. I have compared this type with fe- 
males of several species I collected in the lower 
Amazon. A male, from a series which includes fe- 
males similar to the holotype, was chosen to serve 
as a plesiotype (Ross, 1972: 138) and is the basis for 
figure 20 (below). The task of associating the cor- 
rect male with the female type was simplified by 
the female's distinctive coloration. It has a whitish 
band between each thoracic somite, a feature which 
must have suggested the name, bicingillata. 

Redescription of Enderlein's type (female). — 
Appearance: Moderately small, piceous black 
throughout except for its dark brown head and two 
cream white thoracic bands. Color details (before 
slide preparation): Cranium dark chestnut brown 
throughout blending to piceous laterally and cau- 
dally. dark amber ventrally; without dorsal pattern, 
surface alutaceous; eyes piceous, antennae imi- 
formly medium brown, apical segments lost. Entire 
thorax and abdomen piceous black, blending to ma- 
hogany brown laterally, except for a conspicuous 
cream white band between pro- and mesothorax 
and meso- and metathorax. All legs entirely piceous 
brown or black. Cerci medium brown. All body 




vestiture golden. Body length (on slide) 10.0 mm. 

Discussion. — Males can be readily recognized 
by reference to my 1972 figure (here republished as 
Fig. 20). Characters include the triangular, short, 
fleshy flange of the left tergal process and the very 
long, sclerotic talon of the right tergal process. The 
species may prove to be an Amazonian "weed," ap- 
parently having been extensively moved about in 
river commerce. A specimen from Tobago appears 
to be this species. 

Records (all CAS except otherwise indicat- 
ed). — Brazil: Manaus, on tree bark in city park (E. 
S. Ross); Coragao near Macapa (E. S. Ross); Santa 
Isabel, Rio Arguiya, Goias, at light in forest (B. 
Malkin); Uaupes, Rio Negro, on bark in plantation 
(D. Lacombe); Mun. Boa Vista, Fazenda do Calio- 
clo (Evangelista); Rio Preto "zw Boqueraou Sta 
Rita" (1 male, Vienna Mus.) Guyana: Blairmont. 
Oct. 1923 (E X. Williams). Tobago: Pidgeon Point 
(J. S. Edgerly), probably introduced in commerce. 

Pararhagadochir christae Ross 

(Figure 21) 



Pararhagadochir christae Ross, 1972:135. 
Pararhagadochir cristae Szumik (sic), 
cladogram). 



998:141 (in 



Figure 20. Pararhagadochir hiciiigillata (Enderlein). 
Type locality: Para [Beiem], Brazil. Figure after Ross. 
1972. 



Holotype. — Male, on slide, CAS. Data. — Bra- 
zil: Belem, matured in culture 679, 18-IV-64 (E. S. 
Ross). 

Name basis. — Named after the late Christa Sat- 
tler of the Max-Planck-Institut fur Limnologie who 
sent me specimens and cultures from Belem. 

Redescription (males). — Antennae especially 
long due to elongate segments; segment 1 dark 
brown, 2-3 pale amber blending distad to light 
brown, 3-24 especially long, with wavy setae. Sub- 
mentum heavily sclerotized, with sides and anterior 
margins inflexed. Papilla of hind basitarsus a small, 
flat membranous spot. Wings narrow; hyaline 
stripes narrow, margins sharply defined; in 
forewing the fork of MA is within basal third, only 
one cross-vein between RP and MAj+i, none be- 
hind MA, +2- Terminalia with 10 LP relati\'ely short, 
its flange sclerotized but exceptionally small; api- 
cal talon of 10 RP and its subtended lobe very 
small, both directed downward; medial flap (MF) 
sclerotic, strongly arched, spiculations in cleft 
membrane very fine, almost invisible; epiproct 



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Figure 21. Pararhagadochir chrisTae Ross. Type locali- 
ty: Belem, Brazil. Figure after Ross, 1972. 

sclerite (EP) very large, flared caudad; caudal mar- 
gin of left paraproct (LPPT) without spine, lobe, or 
microspiculation; left cercus lobe relatively small, 
somewhat hooked toward base of cercus. 

Discussion. — In 1964 I found this very distinct 
species abundant on tree bark in park-preserved 
patches of original forest within the city of Belem. 
Pararhagadochir biclngillata, found in the same 
habitats, is readily distinguished by male terminalia 
characters and even more easily by the number of 
pale thoracic bands in adult females; one in P. 
christae, two in P. biclngillata. 

I have collected this species, or a close relative, 
under stones in a low thorn forest in northeastern 
Brazil near Minas do Uranio, Ceara. 

Pararhagadochir balteata Ross 

(Figure 22) 

Pararhagadochir balteata Ross, 1972:133. — Szumik, 
1998:141 (in cladogram). 

Holotype. — Male, on slide, CAS. Data. — Bra- 
zil: Sao Paulo, matured in culture C-706, 27-X-64 
(E. S. Ross). 

Redescription (Males). — Antennae especially 
long, due to elongate segments; uniformly light 
brown to apex, setae in basal half very long. Sub- 
mentum heavily sclerotized, its sides and anterior 
margins inflexed. Papilla of hind basitarsus very 
small, scarcely inflated. Wings narrow, venation 
strong, pigment borders of RBS bright purple; in 




Figure 22. Pararhagadochir balteata Ross. Type locali- 
ty: Sao Paulo, Brazil. Figure after Ross, 1972. 

forewing, fork of MA is just within basal half; three 
cross-veins between RBS and RP, one between base 
of RP and MA, +2, one between MA and MP one 
between base of MA3+4 and MP; hyaline stripes 
narrow, borders sharply defined. Terminalia similar 
to that of P. trinitatis. Apical talon of 10 RP large, 
down-curved, subtended ventral lobe large, mem- 
branous. Medial flap (MF) narrow, not strongly 
arched; spiculations in cleft membrane extensive 
but very fine. Epiproct sclerite (EP) small, narrow 
at base, flared caudad. Caudal margin of left para- 
proct (LPPT) without a spine, lobe or microspicu- 
lations. 

Discussion. — This species is most readily dis- 
tinguished by the pale bands between all body 
somites of adult females. Colonies are conspicuous 
on the bark of trees due to the white, uncovered sur- 
face of gallery silk. 

For convenience, my original figure of the ter- 
minalia of the holotype is here republished as Fig. 

22. 

Additional record. — Brazil: Usina Ester, near 
Cosmopolis, Sao Paulo (CAS) (E. S. Ross). 

Pararhagadochir tenuis (Enderlein) 

(Figure 23) 

Embia tenius^nAer\t\n, 1909:186.— Krauss, 1911:69.— 
Navas, 1918:103. 



ROSS: EMBIA. BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



53 



Rhagadochir tenuis (Enderlein), Enderlein, 1912:60. 
Pararhagadochir tenuis (Enderlein). Davis. 1940a: 



\i 



-Ross, 1944:431. 



Lectotype by present designation. — Male on 
slide deposited in Polska Academia Nauk, Warsaw. 
Labels. — "Boiivien Prov Sara, Steinbach S," on 
green card, "Embia tenuis Enderl. Male Type det. 
Dr. Enderlein," "Type" on red card. 

Locality interpretation. — Prov. del Sara appears 
to be the old name of Prov. Gutierrez, Dept. Santa 
Cruz, about 100 km NW of Santa Cruz de la Sier- 
ra. 

Description of lectotype. — Appearance: Rather 
small, slender, wings relatively large; color uni- 
formly medium brown. Color details (dry on pin): 
Cranium medium chestnut brown, lacking pattern. 
Eyes black. Antennae concolorous with cranium, 
color uniform from base to apex (20 segments pres- 
ent). Mandibles pale amber, remainder of specimen 
varied shades of medium brown, almost as dark as 
cranium. Lacking a pale band between pro- and 



pterothorax. Legs with paler femur-tibial joint. Di- 
mensions (on slide): Body length 9.0 mm; forewing 
length 6.0 mm, breadth 1.5 mm. 

Anatomical distinctions. — As figured. Most 
notable is the head with very large, coarsely- 
faceted, bulging eyes separated by little more than 
an eye-width. Also, the strongly caudal ly-conver- 
gent short sides, only one eye-length long. The 
flange of the left tergal process is broad, rather 
evenly margined and transparent apically. The mi- 
crospiculate nodule of the left paraproct is narrow, 
abruptly rounded. 

Females. — Uniformly chestnut brown with a 
conspicuous cream white band between each tho- 
racic somite. Femur-tibial joints of mid and hind 
legs whitish; hind basitarsal mid-papilla present 
(absent in males). 

Records. — Bolivia: Santa Cruz, males at light 
(G. Pinckert); Santa Cruz (C-768), large cultured 
series; stock from bark flakes of garden trees in 
town (E. S. Ross). My record (1944:431) of this 




TERMINALIA 

VENTRAL 

Figure 23. Pararhagadochir tenuis (Enderlein). Type locality: Santa Cruz region, Bolivia. Figure based on lectotype. 



54 



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species from Rurrenbaque, Bolivia (USNM) is 
questionable in view of distinctions in the head and 
left tergal process and distinct habitat. 

Discussion. — Although closely related to P. 
birabeni, P. tenuis males are readily distinguished 
by the absence of a whitish band between the pro- 
and ptero thorax. The head and eye characters are 
also useful, being invariable in my large series. The 
head of P. birabeni is oval with less convergent 
sides and small eyes separated by at least four eye- 
widths. 

A male of a possible new species, more related 
to P. birabeni than to P. tenuis, was collected by 
Borys Malkin in Bolivia at Coroicom, Yungas Prov. 
It has large wings and apparently no pale thoracic 
bands (not recorded before slide preparation), head 
and eyes similar to P. birabeni. The talon of the left 
tergal process is evenly curved, not abruptly as in P. 
tenuis and P. birabeni, and the paraproct lobe is 
larger and very broadly rounded. 

Another discrepant male, collected by me at 
Angostura, 70 km SW of Santa Cruz, Bolivia, has 
many P. birabeni characters, especially in its oval 
cranium (with very strong pattern) and small, wide- 
spaced eyes. However, all membranous areas of the 
thorax are heavily tinged with dark purple, there- 
fore, as in P. tenuis, there are no whitish intersomi- 
tal bands on the thorax, as is characteristic of P. 
birabeni. 

Pararhagadochir birabeni (Navas) 

(Figure 24) 

Embia birabeni Navas, 1918:105.— Davis, I940b:352 
(Embolyntha?). 

Pararliagadochir birabeni fNavas). Ross. 1944:430. — 
Szumik, 1998:141 (in cladogram). 

Embia piquetana Navas, 1919:25. — Davis, 1940b: 
352. — Ross, 1944:497 (as unrecognizable). New syn- 
onym. 

Lectotype, by present designation. — Male, on 
slide. La Plata Museum, Argentina. Data. — Argen- 
tina: "Unquillo (Cordoba) dr. Max Biraben." De- 
tails of labels in Ross 1944:431. 

Discussion. — I have also studied a badly dam- 
aged cotype male with lectotype data, labeled 
"Typus" in the Navas collection, Barcelona. Be- 
cause of its better condition and deposition in an 
Argentine museum, the above cited cot>'pe was 
chosen to serve as lectotj'pe. 



Also studied in the Navas Collection when it 
was in Zaragoza (1960), was "Typus" o^ Embia pi- 
quetana Navas. It is a mature female labeled in 
Navas' hand: "Santa Fe (Argentina) 22-1-18" 
"Embia piquetana Female Nav.:Navas SJ det" 
"Typus" (pink paper), I found no difference be- 
tween this female and those associated (by rearing) 
with typical males of P. birabeni. Therefore, unless 
the Santa Fe population should later prove to repre- 
sent a distinct species or race, the name E. pique- 
tana should be regarded as a synonym of P. 
birabeni. 

Females. — Almost identical in appearance to 
those of P. tenuis in spite of the fact that P. tenuis 
males are unicolorous (without pale thoracic 
bands). 

Biology. — Pararhagadochir birabeni is found 
in a wide variety of Argentine habitats: in bark 
flakes and under stones in seasonally dry arid zones 
characterized by thorn-scrub and cactus. It is also 
abundant, at least as high as 2000 m elevation, 
under stones on barren grassy slopes in the Volcan 
region of Jujuy. 

Records. — I have cultured several large series 
from Argentina's Rio Quarto, Cordoba in the south 
to just beyond its Bolivian frontier in the north. 
Within this range, especially in the Andes, there is 
much variation, but this may be clinal. Therefore, 
one should hesitate to apply formal nomenclature 
to these locality-associated variants. Geographic 
vernacular names should serve. The most interest- 
ing variations involve the length and size of the 
wings. Such variation is cited in the following 
records (C-numbers = culture numbers). 

Records. — Cordoba: 45 mi. N Rio Quarto (C- 
763), normal wings; Cosquin, Sierra de Cordoba 
(IM, Cornell U). Catamarca: Frios, normal wings. 
Salta: Campo Quijane (P. Wygodzinsky), normal 
wings: Yatasto, near Metan (C-754), normal wings; 
Cabeza de Buey (C-756), normal wings; 20 mi. S. 
Rio de la Frontera, normal wings; 5 and 50 km S. 
Oran (C-759, 760), normal wings; 3 mi. S Salta, 
most specimens with miniature wings; a few with 
wings normal. Jujuy: 5 mi. NW Jujuy, 1000 m, all 
with miniature wings; Arroyo Yuto (C-762), normal 
wings: CTA-9 at Salta border (C-765), normal 
wings: 5 km S San Pedro (C-763), wings normal; 3 
mi. S Volcan, 2000 m, wings half-size. Bolivia: 
Yacuiba (airport at S port of entry) (C-766), small 



ROSS: EMBIA. BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



55 




flange -X^^ talon 

10 RP -~ ' LEFT TERGAL PROCESS 

Figure 24. Pararhgadochir birabeni (Navas). Type locality: Unquillo, Cordoba, Argentina. Figure based on lectotype. 



TERMINALIA 

VENTRAL 



males with very small wings. Unless otherwise in- 
dicated all collected by me and deposited in CAS. 

Pararhagadochir confusa Ross 

Pararhagadochir confusa Ross, 1944:428; Szumik, 
1998:141 (in cladogram). 

Holotype. — Male, on slide, MCZ. Data. — Para- 
guay: Villarica, March (F. Schade). 

The following is a description of an adult fe- 
male from Buenos Aires. 

Description. — Appearance: Moderately large, 
body length 12.0 mm. Blackish brown throughout 
with a very broad cream white band between each 
thoracic somite. Color details (in alcohol): Cranium 



uniformly dark brown; antennae, mouthparts and 
venter of craniuin chestnut brown. Cervical scle- 
rites dark amber, ventral cervical membranes 
tinged with dark purple. Thoracic and abdominal 
tergites dark brown except for pale amber acroter- 
gites which, with the extensive cream white mem- 
branes, form a pale broad band between each tho- 
racic somite. Thoracic acrotergites shades of yello\\' 
tan; ventral sclerites, pleura and associated mem- 
branes dark purple brown. Legs dark brown except 
for chestnut coxae and trocanters of mid and hind 
legs. Abdominal tergites dark brown; except eighth 
and ninth which are translucent pale amber; all ster- 
nites and membranous areas dark purple. Para- 
procts and cerci dark brown, including joint 



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OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES 



membranes. Hind basitarsus with a large medial 
papilla. 

Discussion. — Pararhagadochir confusa is very 
closely related to P. tenuis and P. birabeni but 
males are distinguished by the less abruptly curved 
talon of the left tergal process (10 LP) and the more 
regular shape of its outer flange. They also average 
larger size, much darker color and have a broader, 
more quadrate cranium. Their eyes are larger than 
those of P. birabeni but much smaller than those of 
tenuis. 

Pararhagadochir confusa apparently is wide- 
spread in SE Paraguay, eastern Argentina and SE 
Brazil and probably is spread in human commerce. 
I found it common on the bark of shade trees in 
Corrientes, Argentina and Robin Leech provided a 
culture from tree bark in the city zoo, Buenos Aires. 
Adults in this culture matured during all months of 
the year. In addition, I have a series of males prob- 
ably of the holotype series from the Crampton Col- 
lection. One male in this lot, apparently collected at 
light, is only half the size of P. confusa and has a 
head similar to that of P. birabeni and a distinct left 
cercus lobe. It may represent an eastern population 
of P. birabeni and therefore would indicate that the 
two species can be sympatric. 

Pararhagadochir minuta Ross 
new species 

(Figure 25) 

Holotype. — Male, on slide, MZUSR Data.— 
Brazil: 37 km NE Taua, 425 m, Ceara. Matured in 
culture lO-IV-99 (E. S. Ross). 

Description. — Appearance: Smallest species of 
the genus (body length 7.5 mm), alate, almost black 
except for a white band between pro- and mesotho- 
rax. Color details (in alcohol): Cranium entirely 
black except for its lighter ventral bridge, dorsal 
surface finely alataceous. Eyes dark purple, anten- 
nae exceptionally long; segment (scape) black, 2nd 
brownish black, other segments at first dark amber, 
but grading distad to medium brown; 19-segment- 
ed (complete), flagellar segments bearing excep- 
tionally long setae. Mandibles dark amber except 
for piceous margins and dentation; other mouth- 
parts dark brown except for a brownish black sub- 
mentimi. Thoracic and abdominal sclerites varied 
shades of dark chesmut brown; membranous areas 
pinkish tan except for a white (due to internal fat) 



dorsal band between pro- and mesothorax. Legs 
concolorous with thorax. Sclerotic portions of ter- 
minalia (including segment 9) piceous black, talons 
of 1 LP and 1 RP very dark amber, membranous 
areas pinkish tan; LC] dark brown except for its 
cream outer basal angle, RC, yellow cream except 
for inner margin; joints and tips of discal segments 
white, otherwise pale brown. Dimensions (on 
slide): Body length 7.5 mm; forewing length 3.5 
mm, breadth 0.9 mm. 

Important anatomical features. — As figured. 
Submentum much broader than long, only moder- 
ately sclerotized, anterior margin weak, not inflexed 
(as in congeners); mentum well separated. Wings 
rather small, narrow, hind margin very narrowly ta- 
pered toward base; hyaline stripes narrow, well de- 
fined; no cross-veins; longitudinal veins MA, MP 
and Cu traceable only by their setae. Marginal lines 
of radial blood sinus (RCB) gray white instead of 
the usual pink. Hind basitarsus very short; medial 
papilla small, obscure; plantar setae large, basals 
which are slanted distad are broad; distals slanted 
basad are slender. Terminalia, as figured, mi- 
crospiculate depression in cleft of 10 very large. 
Talon of 10 LP evenly curved and tapered; its 
flange short, basally sclerotic, its small apex white, 
unsclerotized. Dorsal portion of 10 RP irregular, 
truncate; its talon short, basally sclerotic, its small 
apex white, unsclerotized. Dorsal portion of 10 RP 
irregular, truncate; its talon short, narrow, directed 
ventrad. Epiproct sclerite (EP) prominent, very 
long, flared caudad from a long narrow base. Left 
paraproct (LPPT) very large closely appressed to 
HP, inner caudal angle not produced as a spine, or 
as a microsetose nodule. Lobe of LCi very large, 
margin a perfect semicircle, very densely and fine- 
ly echinulate. 

Allotype. — Female, in alcohol, MZUSP, from 
holotype 's culture. 

Description. — Appearance: Generally very 
dark brown except for a pale band between each 
thoracic somite. Antennae and cerci tan. Color de- 
tails: Cranium dark on vertex, grading to amber 
yellow toward clypeus; vertex pattern very faint. 
Eyes concolorous with cranium. All antennal seg- 
ments tan, joints cream, 24-segmented. All dorsal 
body sclerites and legs (except for cream femur-tib- 
ial joints) glossy dark chestnut brown; most mem- 
branous areas brick red, almost concolorous with 
sclerites. Acrotergite 1 and adjacent sclerites clear 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



57 




Figure 25. Pararhagadochir minuta Ross, n. sp. Type locality; NE Brazil: 37 km N Taua, 425 m, Ceara. 



pale amber; these combined with white surround- 
ings (due to color of interior fat) create a broad pro- 
mesothoracic band. A similarly-produced but nar- 
rower band is present between meso-and metatho- 
rax. All ventral body sclerites, except meso- and 
metastema, which are mottled with rust brown, are 
clear amber. Those of prothorax, cervix, and para- 
genital stemites dark chestnut brown. Para-anal 
portions of paraprocts white due to fat visible 
through their translucent derm. Cerci very pale tan 
except for cream base of basal segments. Body 
length 10.0 mm. 

Paratypes and parallotypes. — Adults from holo- 
type's culture deposited in CAS, USNM, MNRJ 
and BMNH. 

Additional records (all NE Brazil). — Paraiba: 
15 km SE Patos; Microware station above Sao 
Bentinao. Piaui: 15 km N Sao Raimundo Nonato, 
500 m; 24 km SW Picos. Ceara: Mina do Uranio. 
Bahia: 20 km SW Casa Nova (northern Bahia). 



Specimens were collected in galleries at the 
edges of small stones and in adjacent leaf litter be- 
neath low trees in usually dry habitats. Fortunately, 
during March 1999 there had been abundant rain- 
fall, otherwise, embiids wouldn't have been present 
in new surface galleries. In cultures adults of both 
sexes matured throughout 1999 and increasingly- 
large cultures were maintained during 2000. 

Discussion. — Pararhagadochir minuta is per- 
haps at least subgenerically distinct from other 
Pararhagadochir, as suggested by its very small 
size; the broad, weakly-sclerotized submentum 
with a non-inflexed anterior margin (a sclerotic, 
shield-like submentum is one of the characteristics 
of all other species of Pararhagadochir): the basal- 
ly-tapered, small wings with weak veins and no 
cross-veins; and the exceptionally large, evenly- 
rounded left crcus lobe. 



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Genus Litosembia Ross 

new genus 

Type species. — Litosembia oligembiodes Ross, 
new species. 

Distribution. — South America: Lower Amazon 
(one record). 

Name basis. — Greek litos (plain, simple) in ref- 
erence to the small, simple appearance. 

Diagnosis. — Males very small (body length av- 
eraging 6.5 mm), alate; pale straw yellow, cranium 
darker with black eyes. Cranium circular in outline; 
eyes very large, inflated, interspace one eye-width, 
facets large; antennae unicolorous, 16-segmented; 
mandibles oligotomoid, incisor teeth well spaced; 
submentum shield-like, sclerotic, anterior and later- 
al margins inflexed, anterior angles projected for- 
ward. Hyaline stripes of wings broad, margins ir- 
regular; MA branches, MP and Cuia unsclerotized; 
cross-veins behind RP almost obsolete. Hind ba- 
sitarsus without a second (medial) papilla; plantar 
setae large. Terminalia weakly sclerotized, sclerite 
margins faint; cleft of tenth tergite broad to basal 
margin. Left tergal process (10 LP) very short, 
broad, not projected beyond caudal margin of para- 
proct; inner margin sclerotized and produced cau- 
dad as a tiny, spine (talon); outer appendix (flange) 
broad, its outer margin indistinct. Basomedial and 
outer margins of right hemitergite (10 R) indistinct, 
mediocaudal margin strong; apex rugosely round- 
ed, but not developed as a distinct process. Medial 
flap (MF) not developed; anterior extremity bearing 
a small, microspiculate nodule. Epiproct sclerite 
scarcely discernable. Ninth sternite (H) broad, mar- 
gins indistinct; its lobe (HP) broadly rounded at 
apex, its right side narrowly sclerotic, almost fused 
to slender right paraproct (RPPT). Left paraproct 
(LPPT) fused basally to H; bearing a non-spiculate, 
elongate nodule on inner-caudal angle. Basal seg- 
ments of cerci with outer side membranous, inner 
side sclerotized; left bearing a large distal lobe hav- 
ing numerous, coarse echinulations and it is arced 
basad. 

Females: With distinct coloration but no gener- 
ic-level anatomical characters except for the pres- 
ence of a second hind basitarsal papilla. 

Discussion. — Litosembia is one of the most 
distinct genera of American Embiidae. Because of 
its small size and coloration, it appears to be a ter- 
atembiid but there is no question that it is a scelem- 



biine embiid. I am inclined to relate it to the genus 
Pavarhagadochir. Its most distinctive characters 
include the broad, short, left process (10 LP) with a 
minute inner spine; the subobsolete right process 
(10 RP) and medial flap (MF) (except for its 
minutely spiculate nodule). The apparent absence 
of a second hind basitarsal papilla in males is also 
diagnostic. Reduction of structures apparently cor- 
relates with exceptionally small size — it is the 
smallest known species of Embiidae. 

Only one species has been discovered to date 
but others should be collected in the future. 

Litosembia oligembioides Ross 
new species 

(Figure 26) 

Holotype. — Male, on slide, MNRL Type 
data. — Brazil; 5 km S. Belem, Para. Matured in 
culture May 1, 1975 (E. S. Ross). 

Description. — Appearance: Small, pale with 
mottled, subdermal rust red and yellow areas; eyes 
contrastingly dark. Color details (in alcohol): Cra- 
nium basically golden yellow but clouded subder- 
mally with rust red except for clear vertex pattern. 
Eyes dark lavender. Antennal segments 1 and 2 yel- 
low amber, segments 3-5 pale tan, all others to 
apex (segment 15) medium brown with pink joint 
membranes, apical segments as dark as subapicals. 
Mouthparts concolorous with antennae except for 
dark golden amber mandibles and submentum. Cer- 
vical area and anterior half of prothorax pale amber 
becoming whitish caudad and on coxal membranes. 
Pterothorax paler, especially due to extensive whit- 
ish, weakly sclerotized areas forming a distinct 
white band between meso- and metathorax; various 
promontories tinged with rust red. Wing bands pale 
brown; hyaline stripes broad with irregular mar- 
gins; RBS lines very pale pink. Forelegs rust amber 
except for medial area of femur which blends to 
gray white. Mid- and hind legs transparent gray 
white except for femora which are broadly banded 
with rust amber basally and apically (similar to pro- 
femora). Abdomen pale lemon-yellow ventrally; 
distinctly banded dorsally due to rusty, subcutane- 
ous pigment; terminalia pale, transparent tan except 
for medium brown inner margins of basal segments 
of cerci and entire apical segments. Dimensions (on 
slide): Body length 6.5 mm; forewing length 4.75 
mm, breadth 1 . 1 mm. 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



59 



Important integumental characters: As figured 
and cited in the generic diagnosis and discussion. 

Allotype. — Female in alcohol with same data 
and disposition as holotype except for maturity 
date, October, 1975 (from same culture). 

Description. — Appearance: Small, head golden, 
body rust brown with distinct cream white bands. 
Color details (in alcohol): Cranium bright yellow 
gold with very faint rust pattern lines. Eyes small, 
blackish. Antennal segments 1 and 2 yellowish, 3 
gray white, 4-6 blending from pale tan to light 
brown, 7-15 dark chestnut brown with pink mem- 
branes, 16 (the apical segment) abruptly pale tan 
with whitish apex. Mouthparts pale amber except 
for apices of palpi which become light brown. Cer- 
vical membranes pale pink. Pronotum clear, pale 
amber blending caudad to pale cream; coxal mem- 
branes white. Acrotergite 1 transparent tan, adjacent 
membranes dark rust red. Mesothorax dark rust red 
except for caudal blend to transparent white which, 
with the white acrotergite 2 and its adjacent mem- 
branes, forms a broad, whitish, meso-metathoracic 
band; metathorax largely rust red except for white, 



subcutaneous caudal areas visible through derm. 
Forelegs rusty, mahogany brown except for coxae, 
medial area of femora and distal tarsal segment, 
which are cream white. Mid and hind legs cream 
white throughout except for rust red basal and api- 
cal pigmentation of femora. Cerci rust tan through- 
out (including joint membranes) except for white 
extreme apices of distal segments. Dimensions (in 
alcohol): Body length 8.0 mm. 

Anatomical characters: The very slender propor- 
tions and the presence of a second hind basitarsal 
papilla. 

Paratypes and parallotypes. — Thirteen adult 
males, one female; from type culture deposited in 
CAS, USNM and MZUSR 

Biology. — This species has been collected only 
once. Stock for the type culture was secured on dead 
bark of a small, native tree at the edge of semi- 
cleared, secondary forest in the public park on the 
southern outskirts of Belem, not far from the swim- 
ming pool. Silk galleries were beneath orchid roots 
growing on almost bare bark of the dead tree. Be- 
fore the culture died, it produced a small series of 




HIND BASITARSUS 
Figure 26. Litosembia oligemboides Ross, n. gen and n. sp., holotype. Type locality: Belem, Brazil. 



60 



OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES 



males, most of which matured during the months of 
September and October; one as late as December. 
The pale color and large eyes of the male indicates 
that they probably disperse noctumally and should 
be attracted to lights. 

Genus Biguembia Szumik 

Biguembia S,zam\\i, 1998:149. 

Type species. — Biguembia copa Szumik, by 
original designation. 

Distribution. — Argentina (Santiago del Estero) 
and Brazil (Mato Grosso, and the northeast Piaui). 
Probably also in intermediate regions. 

Discussion. — When the main body of this paper 
was written, I had no specimens of this genus. How- 
ever, unexpectedly, two cultures I secured in north- 
eastern Brazil early in 1 999 are producing hundreds 
of adult specimens of both sexes of a new species 
very closely related to those described by Szumik. 
Therefore, it can serve as the basis for the following 
amplified generic description. 

Generic description. — Males very large, body 
length averaging 15.0 mm. alate, varied shades of 
brown throughout. Cranium about as broad as long; 
ventral bridge short, weakly sclerotized; eyes large; 
antennae very long, 37-segmented (number proba- 
bly variable), unicolorous to apex. Mandibles short, 
outer margins evenly arcuate; apical dentation large, 
well spaced, without proxidental cusps. Submentum 
much broader than long, weakly sclerotized, side 
and anterior margins not inflexed, setae evenly 
spaced; mentum well defined. Wing venation vari- 
able (even in a single specimen) but at least in B. 
midtivenosa, MA always is multibranched (instead 
of only two-branched as in most other Embiidae). 
Rarely MP is also branched. Hind basitarsus with a 
conspicuous medial papilla; plantar setae dense, al- 
most uniform in shape and length. Terminalia with 
tenth tergite broadly cleft, without a microspiculate 
depression in cleft membrane. Left hemitergite (10 
L) strongly transverse, its base almost entirely sep- 
arated from basal margin of tenth tergite by a nar- 
row membranous interval, abruptly curbed caudad 
to form the inner margin of the long, sclerotic talon 
of its process (10 LP) which is directed straight cau- 
dad and sharply tapered: to its left, and well-sepa- 
rated, a basally sclerotic flange extends caudad al- 
most equal in length to the talon: apical two-thirds 
of flange is membranous, flexible and laterally com- 



pressed (in alcholic specimens the flange is stiff, 
but twists out of shape during slide preparation). 
Inner basal portion of 10 R, the medial sclerite 
(MS), is extensively depressed and lacks setae, its 
inner margin (MP) is evenly arcuated and continues 
caudad to become the left side of a peculiar rugose, 
setose lobe (Saoimik's "hunch"). This lobe overlays 
a very long, narrow, spine-like process, or talon (10 
RP) directed ventrad (Szumik figures this talon as 
rather short in her species). Epiproct sclerite (EP) 
long, narrow, inconspicuous due to weak sclerotiza- 
tion. Hypandrium (H) very large, its process (HP) 
very broad, arcuate apically. Sclerite of left para- 
proct (LPPT) very narrow, partially extended be- 
neath edge of HP, but not fused to it or to H; termi- 
nated caudad as a rugose, sclerotic blunt lobe; 
LPPT's membranous inner-apical portion setose. 
Basal segment of left cercus (LC,) very large, dor- 
sally depressed; its straight, arcuate, or bilobed 
inner side is sclerotic and very coarsely echinulate. 
Other cercus segments normal, well sclerotized ex- 
cept for membranous outer base of RCj. 

Females. — Very large (body length averaging 
20 mm); varied shades of mottled blackish brown 
except for a pale band between meso- and metatho- 
rax. Cranium basically dark chestnut brown, heavi- 
ly mottled with dark mahogany brown. Antennae 
very long, segments 1-24 medium brown, thence to 
apex (segment 36) grading to gray white. Acroter- 
gite 1 dark brown; acrotergite 2, prescutum and 
caudal margin of mesonotum, mottled pale amber, 
adjacent membranes cream white due to interior 
fat, this ensemble produces a pale intersomital 
band. Femur-tibial joints of mid and hindlegs 
whitish. Venter of body paler than dorsum; pleura 
of abdomen, tan to black not forming pale lateral 
stripes. Eight abdominal stemite pale, microhirsute 
medially, each side with an irregular dark brown 
pattern; ninth stemite uniformly mottled brown, its 
anterior margin partially overlays a deep, golden, 
sclerotic pouch (aperture of accessory gland ?). 
Cerci dark purple brown, including joint mem- 
branes. 



ROSS: EMBIA. BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



61 



Biguembia miiltivenosa Ross 
new species 

(Figure 27) 

Holotype. — Male, on slide, MZUSP. Data. — 
Northeastern Brazil: 24 km SW Picos, Piaui, ma- 
tured in culmre l-VI-99 (E. S. Ross). 

Name basis. — Refers to variable multibranched 
MA and MP wing veins. 

Description. — Appearance: Size, coloration and 



anatomical features as described for the genus. Wings 
unusual in having secondary branching of MA2^3 into 
MAt and MA, and secondary branching of MA4+5 into 
MAj. Terminalia apparently very similar to those of 
Szumik's species, however, she figured 10 RP's talon 
relatively short (or is it foreshortened?). In B. miilti- 
venosa it is extremely long and slender — perhaps the 
longest of any species of the order. Another distinction 
is the bilobed basal segment of the left cercus (inner 
margin straight or convex in Szumik's species). 




HEAD 



Figure 27. Important characters of Biguembia miiltivenosa Ross, n. sp. Wings with unusual venation. In some specimens 
MAij.2 may be simple with only M3+4 forked, or in others the converse; MP rarely is forked. In the insert, 10 R. its nod- 
ule and the long very slender talon, area is figured in profile. Type locality: NE Brazil, 24 km SW Picos, Piaui. 



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Allotype. — Adult female, in alcohol, from ho- 
lotype's culture deposited in MZUSP. 

Description. — As presented in the generic diag- 
nosis. 

Paratypes and parallotypes. — Numerous adults 
from holotype's culture deposited in CAS, USNM, 
MZUSP, MNRJ and BMNH. 

Additional records. — Piaui: East of Nazari do 
Piaui (E. S. Ross) CAS. From a thriving culture in 
my Academy laboratory, first instar nymphs ap- 
peared mid-II-00. 

Discussion. — Future collecting and studies may 
reveal that B. multivenosa and Szumik's species 
may be varied geographical populations of a single 
species. However, amongst hundreds of topotypic 
males of 5. multivenosa. LC, invariably is bilobed 
and MA always is multibranched. Szumik's series 
were too small to test consistency of characters of 
her species. 

Biology. — The type cultures were secured on 
semishaded surfaces of spectacular, reddish sand- 
stone cliffs next to a commercial, spring water, bot- 
tling enterprise. During March each colony com- 
prised an extensive silk mat occupied by scores of 
young nymphs attended by a parent female. No 
adult males were present but these began to mature 
in cultures during late December 1999 increasingly 
so during January 2000. Therefore, the culture cal- 
endar coincides with that in the field, thus indicat- 
ing a single annual brood. The galleries were very 
conspicuous, the silk being white, not coated with 
debris. 

Less conspicuous on the cliffs were colonies of 
Pararhagadochir mimita and those of one or more 
species of Teratembiidae. All species utilized rock 
crevices as refuges from predators and excessive 
heat. 

Later in March 1999 a single culture assumed to 
be B. multivenosa. was secured in a very distinct 
habitat — a dense, low, natural, apparently non-de- 
ciduous forest a short distance east of Nazari do 
Pinhui, Piaui (about 42 km SE of Floriano, about 
84 km W of the type locality of B. multivenosa). 
Here colonies were very rare, being concealed in 
cre\'ices of ver>' hard coarse, dry bark of large 
trees. Third instar nymphs were secured with great 
difficult)'. No adults were encountered in the field 
but the cultures yielded adults late in November, 



1999. Eggs were laid in single-layered masses — 
more than one hundred per mass. The eggs were 
protected by inter-egg, masticated paste placed by 
the parent female. 

Adult females and juveniles are much darker 
than those of the type population due to the fact that 
all intersclerotal membranes are dark lavender 
black, whereas those of Picos females and larvae 
are consistently tan. However, anatomical charac- 
ters of males appear to be identical. I will deter- 
mine taxonomic status by hybridizing cultures. 

Biguembia copo Szumik 

(Figure 28) 
Biguembia copo Szumik, 1998:149. 

Holotype. — Male, IFML. Data. — Argentina: 
Santiago del Estero, Reserva Provincial de Copo, 
X-1989 (J.R Pelotto). 

Discussion. — Szumik illustrated the mentum as 
fused to the submentum (her Mn + Sm). Such fu- 
sion probably doesn't occur and requires confirma- 
tion. She states that the cubitus wing vein is forked 
but this isn't shown in her figure, unless she regards 
the distal half of the cubital blood sinus (CuBS) as 
a vein which is doubtful because the trachea of Cu 
emerges from the sinus well before it reaches the 
wing's margin. She states that R, (my RA or radial 
blood sinus, RBS) lacks longitudinal pigmented 




LEFT TERGAL PROCESS 



HIND BASITARSUS 

VEKTRAi 



Figure 28. Biguembia copo Szumik. Tj'pe locality: Santi- 
ago del Estero, Argentina. Modified Szumik figures. 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



63 



borders; if so, her males are the only specimens of 
the order with such a condition. Her few specimens 
have the normal, single forking of MA characteriz- 
ing most Embiidae. MA in B. multivenosa is addi- 
tionally branched in all examined specimens (about 
100). Therefore, it probably is a good species char- 
acter. The talon of 1 R is figured by her as rela- 
tively short on B. copo and broadly acute in con- 
trast to the much longer, narrowly-tapered condi- 
tion in B. multivenosa. Biguembia copo apparently 
lacks a lobe (her "hunch") above 10 RP s talon and 
LCi has a broadly rounded, not bilobed, inner face. 

Females. — Unknown. 

Paratypes. — Specimens from Argentina: 2 km 
W. Hickman, Ruta Nac. 81, 27-1-95 (Szumik and 
Goloboff), IML. "Collected on wet soil, many 
cross turmels with a lot of spongy web." 

Biguembia cocum Szumik 

(Figure 29) 

Biguembia cocum Szumik, 1998:152. 

Holotype.— Male, MZUSR Data.— Brazil: 
Mato Grosso, Serra do Urucum-Corumba, 30-XI- 
60 (K. Lenko). 




LEFT TERGAL PROCESS 



MIND BASITARSUS VENrRAi 



Discussion. — Biguembia cocum appears to dif- 
fer from B. coco in the shape of 10 RP's talon, the 
presence of a lobe above base of the talon and the 
flat, instead of rounded inner face of LC]. 

Females. — Unknown. 

Paratype. — Five topotypic males, presumably 
in MZUSP and IFML. 

Genus Argocercembia Ross 

new genus 

Type species. — Argocercembia guyana Ross, 
new species, by present designation. 

Distribution. — South America: Amazon Basin 
and Guyanan region. 

Name basis. — Greek argos (white, bright) in 
reference to the almost entirely white cerci of the 
species. 

Diagnosis. — Males small (body length averag- 
ing 7 mm), alate; dark brown to pale tan, surface 
usually with metallic-gold sheen; antennae unicol- 
orous; cerci with apical segments pale; tibial and 
wing bases whitish. Cranium short, eyes generally 
large, inflated; antennae 20-segmented, without 
white apices; mandibles delicate, oligotomoid; sub- 
mentum quadrate, weakly sclerotized, anterior 
margin not infle.xed. Wings with apices of veins 
MA 



1+2' 



Figure 29. Biguembia cocum Szumik. Type locality: 
Serra do Urucum-Corumba, Mato Grosso, Brazil. Modi- 
fied Szumik figures. 



MP and all of Cui reduced to rows of 
setae; apex of RBS curved into subapex of RP; 
cross-veins: 3 RBS - RP, 1 RP - MA2+3, no 
MA2+3- MA4+5 and 1 MA - MP; cross-veins never 
white when crossing hyaline stripes. Hind basitarsi 
with second papilla very small. Terminalia broadly 
cleft to basal margin. Left hemitergite (10 L) in- 
flexed along outer and inner margins, but not along 
its weak caudal margin; mesal angle gradually nar- 
rowed to form a simple, acutely and narrowly-ta- 
pered process (10 LP) which lacks an outer flange. 
Right hemitergite (10 R) equilaterally triangulate, 
inner-basal margin weak; process (10 RP) very 
short, complex, composed of a short talon subtend- 
ed by a small, ventral, membranous lobe. Medial 
flap (MF) prominent, projected almost straight for- 
ward, surface conspicuously strigose; membrane at 
apex complexly wrinkled, microdimpled but very 
weakly microspiculated. Epiproct sclerite (EP) 
large but not extending beyond apex of 1 LP. Left 
paraproct (LPPT) with caudal margin smoothly, 
broadly arcuated; extremities not sharply produced; 
ventro-caudal surface densely microspiculated. 



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Basal segment of left cercus short, apical third de- 
sclerotized; almost entire inner side forming a lobe, 
echinulations confined to inner edge of lobe; lobe 
strongly dorso-ventrally depressed. All other cercus 
segments desclerotized, appearing white in life, but 
membranous in cleared preparations. 

Females: Without noteworthy generic charac- 
ters except the slender body, the golden surface 
sheen, the lack of white apical antennal segments, 
the pale meso-metathoracic band, the whitish tibial 
bases and apical cercus segments. 

Discussion. — The presence in my collection of 
four new species possessing the above characters 
strengthens this generic concept. Actually, it is not 
very closely related to any other known genus ex- 
cept Xiphosembia n. gen., from which it differs in 
many characters, e.g., shorter and broader-based 10 
LP: scarcely-produced trifid 10 RP: and short, de- 
pressed basal segment of the left cercus. 

I collected three additional species in the fol- 
lowing Brazilian localities: Vila Amazona, Amapa 
(semicleared rainforest): Mata da Pirelli, near 
Belem, Para (virgin rainforest); and 20 km N 
Caracarai, Roraima (stones in seasonally-dry grass- 
land). Probably males of some, or all, of these 
species may be attracted to lights. 

Argocercembia guy ana Ross 
new species 

(Figure 30) 

Holotype. — Male, on slide, CAS. Data. — 
Guyana: Atkinson Airport. Georgetown 31-V-64 
(E. S. Ross). 

Description. — Appearance: Rather small, alate, 
light chestnut brown except for tan dorsum of 
pterothorax and partly to entirely white apical seg- 
ments of cerci. Color details (in alcohol): Cranium 
chestnut brown with faint vertex pattern, sheen me- 
tallic. Eyes lavender black. Antennae chestnut 
brown to apex except for yellow segments 2-4 (20- 
segmented). Mouthparts \aried shades of brown. 
Prothoracic sclerites chestnut brown with golden 
sheen; membranes tinged with rust brown. 
Pterothorax shades of yellow tan with darker 
promontories; ventral sclerites slightly darker. Legs 
concolorous with thorax except for whitish tibial 
bases and femoral apices. Wings light brown except 
for whitish bases and broad hyaline stripes. Ab- 
domen rust brown dorsally, tan ventrally; termina- 



lia concolorous except for whitish apical segments 
of cerci. Dimensions (on slide): body length 7.5 
mm; forewing length 5.1 mm, breadth 1.1 mm. 

Important integumental characters. — Until the 
three additional congeners are described, males of 
A. giiyana can be identified by reference to figured 
generic characters and locality. 

Allotype. — Female, in alcohol, with holotype 
data and disposition (from same culture). 

Description. — Appearance: Small, slender, 
chestnut brown with whitish band between meso- 
and metathorax. all appendages dark to apex. Color 
details (in alcohol): Cranium golden brown, paler at 
vertex due to brain color within, clypeus darker. 
Antennae and mouthparts as in males. Prothoracic 
and cervical sclerites dark mahogany brown with a 
green gold metallic sheen; all membranes dark pur- 
ple. Meso- and metathoracic sclerites a tone paler 
than those of prothorax, sheen also metallic; mem- 
branes purple except between somites where 
whitish internal tissues produce a whitish inter- 
somital band. Legs concolorous with thorax except 
for whitish femur-tibial joints. Abdomen concolor- 
ous with thorax with golden sheen; basal segments 
of cerci lavender-black with pale spot at base of 
each trichobothrium, apical segments tan. Body 
length: 9.0 mm. 

Important integumental characters: Those of 
specific importance will be described when addi- 
tional new species in my collection are made 
known. Second hind basitarsal papilla inconspicu- 
ous. 

Paratypes and parallotypes. — 1 1 males and 9 
females deposited in CAS, USNM, BMNH and 
MZUSP All from type culture, matured during Jan- 
uary-May and October-December. 

Discussion. — The type culture was collected in 
bark crevices of small shade trees adjacent to the 
airport waiting room. Argocercembia guyana is 
most closely related to a new species from the Boa 
Vista. Roraima region of Brazil, difl^ering primarily 
in size and coloration. 

Genus Aphanembia Ross 

new genus 

Type species. — Aphanembia obsciira Ross, new 
species, by present designation. 

Name basis. — Greek aphanes, (unseen, invisi- 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



65 




TERMINAUA 

VENTRAL 
Figure 30. Argocercembia giiyana Ross, n. gen and n. sp., holotype. Type locality: Georgetown, Guyana. 



ble) in reference to hidden colonies under bark 
flakes. 

Distribution. — South America: Amazon Basin. 

Diagnosis. — Males: Rather small (body length 
averaging 8 mm), alate; body light tan, head and 
terminalia cinnamon brown. Cranium rectangulate- 
oval, or circular in outline; eyes rather large, inflat- 
ed; antennae without pale distal segments, 14-16 
segmented; submentum transversely rectangulate, 
anterior margin weak; mandibles flat, thin, short, 
apical teeth well-spaced, sharply acute. Hind ba- 
sitarsus short, with two ventral papillae. Tenth ter- 
gite very broadly cleft to basal margin. Left 
hemitergite (10 L) triangulate; its process (10 LP) 
continuous with sclerotic, ridged, inner margin, of 
10 L and represented as an arcuate narrowly-ta- 
pered inner talon and a lower, narrowly-acute, 
much shorter, translucent outer flange. Right 



hemitergite (10 R) with inner margin straight, outer 
margin sinuous with a small membranous interrup- 
tion at outer base of process; right process (10 RP) 
continuous with acutely narrowed sclerotic apex of 
10 R and formed as a striate downwardly-directed 
talon with a tiny nodule present ventrally beneath 
its base. Medial flap (MF) a sclerotized, strigose, 
forward-projecting "arm." Epiproct (EP) with its 
sclerite large, very weakly sclerotized. Hypandrium 
process (HP) with a deep, basal, membranous 
emargination on left side; its apex weakly sclero- 
tized. Left paraproct large, transversely fiised to H 
at base; its caudal margin well defined, evenly ar- 
cuate; a large, acute, cone-shaped microspiculate, 
forward-directed lobe is highly characteristic. Left 
cercus lobe with a single, broadly-rounded, coarse- 
ly-echinulated, medial lobe; inner-basal margin of 
basal segment continued beneath lobe. 

Females: With distinctive coloration. Cranium 



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translucent gold, prothorax pale amber; most other 
body sclerites dark mahogany brown. Legs exten- 
sively pale, cerci entirely dark. Without distinctive 
integumental characters except for the exceptional- 
ly circular head form. 

Discussion. — Although its integumental char- 
acters do not appear to be very different from those 
of related genera, such as Gibocerciis, Aphanembia 
has a generically distinct biology. Its colonies are 
very obscure, being concealed under bark flakes of 
large trees and stumps. No extensive galleries ex- 
tend beyond edges of the covering flake. Collection 
of specimens thus often requires random lifting of 
bark flakes. Related genera, such as Gibocerciis, 
make extensive, exposed gallery systems occupied 
by many individuals. Broods of Aphanembia ap- 
pear to be small and thus its species will be rarely 
collected. A male of one species was collected at 
light. 

The most distinctive character of adult males is 
the conical ventral lobe of the left paraproct. Adult 
females are unique in being markedly bicolorous — 
the head and prothorax are gold and amber respec- 
tively in sharp contrast to the otherwise dark brown 
body. 

Component species. — My collection (CAS) 
contains small series of additional species from five 
localities in the Amazon Basin: Peru: Tingo Maria 
region; Tambopata, Madre de Dios; Estiron, Rio 
Ampi Yacu, Dept Loreto. Brazil: 20 km N Manaus; 
Arequemes region, Rondonia. Colombia: near Leti- 
cia (at light). In spite of vast geographic separation, 
these populations may at best represent races of a 
single species with male distinctions occurring in 
cranial and eye form, and the shape of the para- 
proct's conical lobe. 

Aphanembia obscura Ross 
new species 

(Figure 31) 
Holotype. — Male, on slide, CAS. Data. — Peru: 
Yurac Plantation. 67 mi E of Tingo Maria, 14-XIl- 
1954 (E. S. Ross). 

Description. — Appearance: Rather small, alate; 
pale tan except for darker head, terminalia, and 
wings. Color details (in alcohol): Cranium cinna- 
mon brown with faint \ertex pattern. Eyes blackish 
with pale margins. Antennal segments 1-8 pale 
amber, others to apex (segment 16) slightly darker. 
Mandibles clear amber with piceous margins, other 



sclerotic portions of mouthparts chestnut brown. 
Body sclerites and legs pale tan, prothorax neither 
darker nor lighter than other portions of body; all 
membranes cream white; wings pale tan, cross- 
veins white when crossing a hyaline stripe; termi- 
nalia sclerites, including cerci, chestnut brown. Di- 
mensions (on slide): Body length 9.5 mm; forewing 
length 6.7 mm, breadth 1.5 mm. 

Important integumental characters. — As fig- 
ured and described in the generic treatment. Of pos- 
sible specific, or subspecific importance, is the 
elongate, rectangulate-oval cranial outline. Males 
from Manaus have a short, circular cranium with 
exceptionally large eyes. 

Allotype. — Female, in alcohol, with holotype 
data and disposition. 

Description. — Appearance: Cranium, prothorax 
and legs golden to yellow, other sclerotized por- 
tions mahogany brown, all surfaces glossy; anten- 
nal apices and cerci not contrastingly pale. Color 
details (in alcohol): Cranium translucent gold, brain 
and muscles visible through surface, otherwise 
lacking pattern. Antennae medium mahogany 
brown from base to subapex, thence becoming tan 
distad; 1 6-segmented. Labrum yellow; mandibles 
amber; other mouthparts tan, apices of maxillary 
palpi red mahogany brown. Prothoracic and cervi- 
cal sclerites amber; pronotum more brownish; 
acrotergite 1 translucent pale amber; surrounding 
membrances pale, transparent. Sclerites of remain- 
der of thorax and abdomen shades of mahogany 
brown, all membranes cream white, color of ab- 
domen increased by reddish brown subdermal tis- 
sue. Legs largely pale yellow, fore tarsi and hind 
femora more brownish. Cerci dark reddish brown. 
Body length 9.5 mm. 

Paratypes and parallotypes. — A few topotypic 
adults deposited in CAS, USNM and MNP 

Biology. — As described in the generic discus- 
sion. 

Genus Ambonembia Ross 

new genus 

Type species. — Ambonembia incae Ross, new 
species by present designation. 

Distribution. — Peru (central Andean altiplano) 
and yungas of Bolivia. 

Name basis. — Greek ambon (ridge) in refer- 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



67 




SUBMENTUM 



TERMINALIA 

VENTRAL 



Figure 31. Aphanembia obsciira Ross, n. gen and n. sp., holotrype. Type locality; 67 mi. E. of Tinge Maria, Peru. 



ence to elevated carinate surface of left hemiterg- 
ite. 

Diagnosis. — Males: Medium sized (body 
length averaging 12 mm), alate; uniformly dark 
brown except for darker terminalia. Cranium oval, 
eyes small; antennae uniformly brown to apex; 
mandibles with well-spaced, large apical teeth; sub- 
mentum quadrate, sides straight, parallel, anterior 
margin unsclerotized, not inflexed. Hind basitarsus 
with a large, medial papilla. Terminalia's tenth ter- 
gite broadly cleft to base. Left hemitergite (10 L) 
strongly ridged, or diagonally carinate, continuous- 
ly with base of its process (10 LP) which terminates 
with a talon-like inner process and an outer semi- 
translucent flange. Inner margin of 10 R blending 
to membrane, caudally tapered and terminated in a 
small sclerotic hook. Medial flap (MF) obsolete, 
represented only by wrinkles in cleft membrane 
which lacks a microspiculated depression. Epiproct 



sclerite (EP) narrow basally, flared caudad. Left 
paraproct (LPPT) large, darkly sclerotized along 
contact with side of HP; inner caudal angle with a 
small, sclerotic point, this point and caudal margin 
of LPPT not microspiculated. Lobe of left cercus 
(LCj) very large, densely echinulated. Basal seg- 
ment of right cercus extensively membranous on 
outer side. Apical segments of both cerci weakly 
sclerotized. 

Discussion. — Ambonembia is somewhat related 
to Malacosembia. as suggested by its obsolete me- 
dial flap. Males differ in the following characters: 
Ambonembia eyes small, interspace five eye- 
widths. Malacosembia large eyes, interspace one 
eye-width. Left hemitergite (10 L) diagonally cari- 
nate in Ambonembia, flat in Malacosembia: 10 LP 
and 10 RP very distinct, as figured. Epiproct (EP) 
well developed in Ambonembia but not in Mala- 
cosembia, or only membranous with caudal margin 



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of LPPT smooth in Ambonembia, entirely mi- 
crospiculate with inner caudal angle lobed in Mala- 
cosembia, instead of a sharp point. 

The following two very distinct species are as- 
signed to this genus. Future studies, hopefully 
based on knowledge of more new species, may in- 
dicate that the two are not congeneric. Among 
many distinctions, those of the left tergal process 
are most striking. 

Ambonembia incae Ross 
new species 

(Figure 32) 

Holotype. — Male, on slide, CAS. Data. — Peru: 
Sacsahuaman (Inca ruins), near Cuzco. Colonies in 
trail bank crevices in grassy area. Matured in cul- 
ture March, 1983 (E. S. Ross). 

Description. — Appearance: Moderately large 
(body length 12.0 mm), alate; generally dark choc- 
olate brown, including all appendages. Color de- 
tails (in alcohol): Cranium dark chocolate brown; 
vertex pattern lighter brown, indistinct; clypeus and 
interocular area darkest; venter of cranium paler 
due to larger, more distinct pattern; luster dull. Eyes 
as dark as cranium, almost black. Antennal scape 



dark brown; flagellar segments at first golden 
brown, thence becoming darker distad, apicals dark 
brown; twenty-three segments in complete anten- 
nae. Labrum dark brown, clypeo-labral membrane 
dark lavender gray; mandibles dark brown, other 
mouthparts medium brown. Prothorax and its pat- 
tern concolorous with cranium. Pterothorax gener- 
ally chestnut brown with margins and sutural areas 
dark chocolate brown, all membranes lavender 
gray. All legs varied shades of chestnut brown. Ab- 
domen essentially concolorous with pterothorax, 
terminalia darker; basal segments of cerci dark 
brown, distal segments chestnut brown. Dimen- 
sions on slide: Body length 12.0 mm, forewing 
length 7.2 mm, breadth 9.0 mm. 

Anatomical features. — As illustrated. Particu- 
larly significant is the left hemitergite (10 L) which 
is longitudinally elevated and depressed in the 
medio-basal corner. Its process (10 LP) composed 
of a narrow, tapered, sinuous, inner talon and a 
broad poorly defined, somewhat fleshy outer 
flange. Right hemitergite ( 1 R) lacks a medial flap 
or sclerotized inner margin; its process is a sharp, 
tapered talon curled downward, scarcely visible 
from above. Lobe of left cercus very large, bulbous, 
densely, microechinulate. 




Figure 32. Ambonembia incae Ross, n. gen and n.sp., holotype. Type locality: Cuzco, Peru. 



ROSS; EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



69 



Paratypes. — Eight males deposited in CAS, 
USNM and MHNP. 

Female. — No specimens. It is safe to assume 
that females are uniformly brown, including all ap- 
pendages. 

Ambonembia adspersa (Enderlein) 

new combination 

(Figure 33) 

Embia adspersa Enderlein, 1909:185. — Krauss, 
1911:669.— ^Navas, 1918:103. 

Rhagadochir adspersa (Enderlein), Enderlein, 1912:58. 

Pararhagadochir adspersa (Enderlein), Davis, 
1940a: 188.— Ross, 1944:434. (both after Enderlein). 

Type. — Male, on slide, Polska Academia Nauk, 
Warsaw. 

Type labels: "Bolivien Prov. Sara, Steinbach S." 
(green label), ''Embia adspersa Ended. Male Type 
det. Dr. Enderlein." 

Locality interpretation. — Prov. del Sara is the 
old name of Prov. Gutierrez, Dept. Santa Cruz, 
about 100 km NW of Santa Cruz de la Sierra, Bo- 
livia. 

Condition. — Originally on pin, with head, pro- 
thorax and forelegs missing, terminalia smashed on 
slide. I carefully transferred the fragments into bal- 
sam on a slide (coverslip supported). Due to Ender- 
lein's poor slide preparation, the ninth stemite and 
tenth tergite are badly fractured and structures dis- 
torted in position. 

Redescription. — Appearance: Large, wings ex- 
tensive; medium brown. Color details (dry): Head 
and prothorax lost, but according to Enderlein, the 
head is dark brown with black eyes. No mention of 
a distinctive prothoracic coloration was made so it 
is assumed that the thorax was entirely dark chest- 
nut brown laterally and ventrally and pale chestnut 
brown dorsally. Mid and hind legs medium chestnut 
brown with tarsi appearing golden due to color of 
dense setae. Abdomen dark chestnut brown. Wings 
light brown; hyaline stripes broad margins indis- 
tinct; veins very straight, simple; cross-veins incon- 
spicuous; two between RES and RP, one subapical 
between RP and MA, none between MA, +2 and 
MA3+4 and one between stem of MA and base of 
MP. Hind basitarsi with second papilla in distal 
third. Tenth tergite broadly cleft but not quite to 
basal margin. Left hemitergite (10 L) inflexed and 
sclerotic only along outer-basal margin; longitudi- 




TERMINALIA VENTRAl 



Figure 33. Ambonembia adspersa (Enderlein). Type. 
Type locality: Santa Cruz region, Bolivia. 



nally elevated. Base of process (10 LP) constricted 
longitudinally sulcate; expanded distally with an in- 
ner, broad-based talon, curved outward at 90° and 
an outer-ventral, membranous, microspiculate, 
rounded lobe. Right hemitergite (10 R) triangulate, 
inner margin weak; bearing a short, claw-like 
process ( 10 RP), directed downward (this may be a 
stub of a broken-off longer process). Medial flap 
absent. Epiproct sclerite (EP) very narrow basally, 
broadly expanded caudad. Hypandrium (H) evenly 
sclerotized margins weak, more sclerotic across the 
base of its extremely short process (HP) which is 
membranous along its caudal margin. Left para- 
proct (LPPT) very large, triangulate; sclerotic 
along its inner-basal margin which is fiised to the 
hypandrium; extremities of its unlobed caudal mar- 
gin minutely produced the outer one microspicu- 
late. Basal segment of left cercus well sclerotized; 
apical lobe an elongate cone densely micro-echinu- 
late on inner surface. Basal segment of right cercus 
small, desclerotized on all but inner surface. 

Female: Unknown. Coloration probably similar 
to that of males. 

Discussion. — This distinct species is known 
only from its holotype. Placement of the species in 
Pararhagadochir was erroneous on the basis of 
many characters, such as: the soft, not sclerotic 



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submentum; the obsolete medial flap and the ab- 
sence of microspiculations in the membrane of the 
medial cleft. 

Genus Malacosembia Ross 

new genus 

Type species. — Malacosembia tiiciimana Ross, 
n. sp., by present designation. 

Distribution. — Argentina (Tucuman region), 
Bolivia (Santa Cruz region). 

Name basis. — Greek malacos (soft) in reference 
to overall weak, or soft, body integument. 

Diagnosis. — Males: Size medium to small 
(body length 7 to 10 mm), alate; generally medium 
brown to pale amber with slightly darker head and 
terminalia. Head, wings and hind basitarsi as in 
Ochrembia except for antennae which are not 
white-tipped. Terminalia's tenth tergite broadly cleft 
to base. Left hemitergite (10 L) about as long as 
broad, its base extended beneath caudal edge of 
ninth tergite (9); caudal margin weakly sclerotized; 
surface relatively flat. Process (10 LP) broad-based, 
arising from inner-caudal portion of 10 L; with a 
short, talon-like, sclerotic, inner portion and a nar- 
row, flap-like, weakly-sclerotized, outer appendix. 
Inner-basal side of right hemitergite (10 R) weak; 
process ( 1 RP) an abruptly narrowed "talon" pro- 
jected ventrad. Medial flap entirely absent. Epiproct 
(EP) obscure, its sclerite faint, if at all visible. Left 
paraproct (LPPT) with caudal margin almost 
straight, microspiculate on inner half, with a small 
acute point on inner corner and a lesser, blunt point 
on outer comer. Basal segment of left cercus with a 
prominent, echinulate, inner lobe. 

Females: With pale coloration of males, without 
white-tipped antennae. With two hind basitarsal 
ventral papillae. 

Discussion. — Although the left tergal process is 
somewhat similar to that of some species of 
Pararhagadochir, Malacosembia is not closely re- 
lated. It is most closely related to Ochrembia but 
easily distinguished by its bifid, shorter, left tergal 
process; the absence of a globose, finely-echinulate 
nodule on the caudal margin of the left paraproct 
and other terminalia characters. The absence of 
white apical antennal segments may also be a con- 
sistent character in both sexes. 

Component species. — Because M. tuciimana is 
represented by the largest series of both sexes, it is 



selected as the type species of the genus. I also col- 
lected a closely related new species, M. yungae, 
near Santa Cruz, Bolivia. 

Malacosembia tucumana Ross 
new species 

(Figure 34) 

Holotype. — Male, on slide, Instituto Miguel 
Lillo, Argentina. Data. — Argentina: Tucuman, ma- 
tured in culture August 24, 1964 (E. S. Ross). 

Description. — Appearance: Moderately large, 
winged; mottled, medium brown throughout with 
darker head; body weakly sclerotized. Color details 
(alive): Cranium basically pale tan but heavily 
tinged with granular, subcutaneous dark brown, 
forming a defmite dorso-basal pattern; ventrally 
paler. Eyes blackish lavender, darker than cranium. 
Basal segment of antenna dark brown, other seg- 
ments light tan; segments beyond 16 broken off (22 
when complete). Mandibles pale amber with red 
amber inner-apical margins; other mouthparts tan 
with distal palpal segments darkest. Submentum 
and mentum weakly sclerotized; subcutaneously, 
granular rust brown. Remainder of insect, including 
legs and membranes, basically cream yellow to tan 
but all dorsal sclerites appear darker due to granu- 
lar, rust brown tinge; pronotum and propleurae 
darkest; ventral sclerites pale tan with thoracic ster- 
nites evenly light brown; all leg segments uniform- 
ly light brown. Wings pale tan with broad hyaline 
stripes having ragged margins; costal and RBS bor- 
ders pink. Terminalia concolorous with abdomen, 
talon-like portions of tergal processes reddish 
amber; microspiculate caudal lobe of left paraproct 
clear yellow amber. Dimensions (on slide): Body 
length 10.6 mm; forewing length 7.5 mm, breadth 
2.0 mm. 

Important integumental characters. — As fig- 
ured. Of special importance are size, form of tergal 
processes and the shapes of the lobes of the caudal 
margin of the left paraproct. Paratypes with com- 
plete antennae indicate that the segment number is 
22 and that the apical segments are concolorous 
with other flagellar segments. 

Allotype. — Female, in alcohol, with holotype 
data and disposition. 

Description. — Appearance: Medium sized, 
golden brown with cream white intersclerotal 
membranes (superficially resembling females of 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



71 




TERMINALIA ventral 
Figure 34. Malacosembia tucumana Ross, n. gen. and n. sp., holotype. Type locality: Tucuman, Argentina. 



Oligotoma). Color details: Cranium basically gold- 
en but mottled with rust brown, producing a sym- 
metrical vertex pattern. Eyes jet black. Basal anten- 
nal segment pale amber; other segments paler 
amber, distals becoming darker; 20-segmented. All 
dorsal body sclerites concolorous with cranium, ex- 
tensively darkened by subcutaneous rust-brown; in- 
tersclerotal membranes and entire venter transpar- 
ent cream white — revealing internal organs. Legs 
pale amber becoming darker distad. Cerci mottled 
with rust-brown. Body length: 10.5 mm. 

Paratypes and parallotypes. — Large series from 
type culture deposited in CAS, IFML, MZUSP, 
USNM, BMNH. 

Additional records (IFML): Several very dam- 
aged males from Canete, Prov. Tucuman 14-X-1965 
(EB 451). Tucuman, on citrus bark, X- 11-65 (R. 
Goldbach). 



Discussion. — Malacosembia tucumana has a 
close relative in Bolivia and it is expected that ad- 
ditional congeners will be discovered in neighbor- 
ing regions, such as Paraguay. 

Biology. — Malacosembia tucumana, collected 
on the bark of shade trees within the city of Tu- 
cuman, probably occurs on native trees in the im- 
mediate vicinity. Bark flakes and crevices serve as 
retreats. It is assumed that this widespread habit 
within the order applies to other species of the 
genus. The pale color and large eyes indicate that 
dispersal is nocturnal and that males are attracted to 
lights. Males matured in culture 749 from June to 
October, 1964. 

Malacosembia yiingae Ross 
new species 

(Figure 35 ) 
Holotype. — Male, on slide, CAS. Data. — Bo- 



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livia: Montero. near Santa Cruz, matured in culture 
August 14, 1964 (E. S.Ross). 

Name basis. — Yiingas is the Indian name for a 
zone on the east slope of the Andes. 

Description. — Appearance: Similar to Oligo- 
toma humbertiana; meditim sized, winged, golden 
brown throughout with gold head antennae and 
eyes darker. Color details (alive): Cranium brilliant 
gold, lacking pattern, faintly tinged with granular 
rust red, rims of antennal sockets brown; ventrally 
straw yellow. Eyes blackish lavender. Antennae al- 
most unicolorous chocolate brown; basal segment 
darkest, sub-basals lighter (segments beyond 14 are 
broken off)- Mandibles straw yellow with piceous 
apical margins; submentum golden with brown lat- 
eral margins, outer mouthparts golden except for 
chocolate brown palpi. Thoracic sclerites various 
shades of light brown with piceous sutures and 



margins; dorsal and pleural membranes strongly 
rust red; ventral membranes golden due to color of 
fat visible beneath unsclerotized surfaces. All legs 
entirely tan, becoming somewhat darker distad. 
Wing bands light brown; hyaline stripes broad with 
margins dotted by circular pigmented spots at bases 
of macrotrichiae; all wing veins behind RBS, in- 
cluding cross-veins, pink; margins of RBS brighter 
pink, RP brown. Abdomen largely granular, rust 
red; tenth hemitergites dark brown, cleft membrane 
cream; hypandrium and paraprocts light brown; 
cerci rust red except for base and inner margins of 
basal segment of left cercus. Dimensions (on slide): 
Body length 7.5 mm, forewing length 6 mm, 
breadth 1.1 mm. 

Important integumental characters. — As fig- 
ured. Differs from M. tucumana in the more wide- 
ly spaced, narrower forks of the left tergal process 




TERMINALIA 

VENTRAL 

Figure 35. Malacosembia yimgae Ross, n. sp., holotype. Type locality: Montero, near Santa Cruz, Bolivia. 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



73 



(10 LP), the smaller right tergal process (10 RP) 
which is almost concealed from above by the 
rounded caudal margin of 10 R, the weak but dis- 
cemable, broad epiproct sclerite (EP), the less acute 
inner process of the left paraproct (LPPT), and the 
more coarsely echinulate left cercus lobes. In addi- 
tion, M. yungae is much smaller and overall more 
darkly pigmented than M. tiicumana. 

Allotype. — Female, in alcohol, CAS. Data. — 
reared in holotype's culture. 

Description. — Appearance: Small, body length 
7.5 mm; generally pale amber. Color details: Crani- 
um amber yellow almost lacking pattern; translu- 
cent, brain visible between eyes; blackish at tentor- 
ial pits. Eyes black. Antennal segments amber ex- 
cept segments 4 and 5 and distals which are slight- 
ly paler, distal segment, the 20th, elongate, cream 
white at tip. Dorsal body sclerites and all legs var- 
ied shades of translucent amber; all membranous 
areas dark cream, almost concolorous with scle- 
rites; abdominal somites 9 and 10 golden brown, 
their stemites and paraprocts pale amber; entire 
venter of specimen cream white. Cerci, including 
joints, mottled medium brown except for paler tip 
of apical segments. 

Paratypes and parallotypes. — A series of both 
sexes reared in the same culture, deposited in CAS. 

Discussion. — Malacosembia yungae is one of 
the smallest species of South American Embiidae. 
It has the general appearance of the pale species of 
Teratembiidae occurring in the Santa Cruz region 
of Bolivia. A comparison of my figures of its con- 
gener shows close relationships. From M. tii- 
cumana, M. yungae differs, as follows: one-third 
smaller size; overall darker pigmentation; forks of 
10 LP smaller, narrower and more widely spaced; 
talon of 10 RP much smaller and almost obscured 
from above by the caudal margin of 10 R; and the 
epiproct sclerite of M yungae, more apparent than 
that of M tucumana, is unsclerotized. 

Biology. — The type culture (C-777A) included 
a number of unrelated species webbing the under 
surfaces of logs and branches on leaf litter in 
thinned natural forest behind the agriculture college 
at Montero. Conditions prevailing in 1964 may be 
considerably changed today. 



Genus Ochrembia Ross 

new genus 

Type species. — Embia wagneri Navas, 1923, by 
present designation. 

Distribution. — North-central Argentina and 
south-central Bolivia (three records). 

Name basis. — Greek ochros (pale yellow, sal- 
low) in reference to the pale coloration of the type 
species. 

Diagnosis. — Males: Moderately large (body 
length averaging 1 1 mm), alate; almost unicolor- 
ous, pale, yellow tan. Cranium large, broad; eyes 
exceptionally large, inflated, interspace less than 
one eye-width; antennae white-tipped; mandibles 
normal, teeth of left mandible equal in size; sub- 
mentum weakly sclerotized, anterior margin not in- 
flexed. Wings large, broad; hyaline stripes broad, 
their margins irregular; veins not straight, curved 
toward contacts with cross-veins; cross-veins nu- 
merous — 3 between MA and MP. Hind basitarsus 
with two ventral papillae. Tenth tergite broadly cleft 
to basal margin. Left hemitergite (10 L) narrowly 
transverse, caudal margin weak, basal margin in- 
flexed; cleft membrane extended leftward behind 
basal margin; its process (10 LP), an extension of 
inner side of 10 L, is arcuated caudad as a large, 
sclerotic, almost simple process (10 LP), its outer 
appendix, or flange, a tiny sliver, is almost com- 
pletely invisible. Right hemitergite (10 R) with in- 
ner-basal margin obsolete, blending into cleft mem- 
brane; caudal margins sclerotized; process (10 RP) 
abruptly formed as a narrow, outwardly and ven- 
trally-angled spine which is subtended by a small 
membranous lobe. Medial flap (MF) obsolete, rep- 
resented only by a weak membranous wrinkle. 
Epiproct (EP) not prominent; its sclerite faint, but 
broad, almost horizontal (90° to axis of insect); left 
portion (beneath basal arc of 10 LP) inflated, 
densely microspiculate. Hypandrium lobe (HP) 
short, broad, truncate. Left paraproct (LPPT) with a 
prominent, broad, submembranous, caudal lobe 
which is densely microspiculate. Basal segment of 
left cercus with a very large, broadly-rounded, 
densely echinulate inner lobe. Basal segment of 
right cercus almost entirely without sclerotization. 

Females: Unknown. Undoubtedly very pale in 
coloration. 

Discussion. — From the only other South .'Amer- 
ican genera with an obsolete medial flap. Mala- 



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cosembia Ross and Ambonembia Ross, Ochrembia 
is distinct in many characters — particularly in the 
almost simple, instead of conspicuously bifid, left 
tergal process. The genus is probably widespread in 
central and northern Argentina, as well as in south- 
em Bolivia (east of Andes) and western Paraguay. 
Specimens from three widely scattered regions are 
remarkably similar. Those (CAS) from near Camiri, 
southcentral Bolivia have only a slightly distinct 
left cercus lobe. Pale color and large eyes suggests 
that the males disperse at night and that specimens 
collected to date were attracted to lights. Colonies 
of this genus should occur beneath coarse bark 
flakes and in crevices of tree trunks, and possibly 
beneath stones. 

The poorly described type species is re- 
described, from its holotype, as follows: 

Ochrembia wagneri (Navas) 
new combination 

(Figure 36) 

Embia ivagwer; Navas, 1924:13, fig. 3. 
Embohmtha wagneri (Navas), Davis, I940b:351, figs. 
384l— Ross. 1944:413. 

Holotype. — Male, on slide, deposited in the 
Museum National d'Histoire Naturelle, Paris. La- 
bels. — "Embia Wagneri Nav. P. Navas S. J. det" 
(yellow) "Type" (printed, red), "Museum Paris 
Chaco de Santiago del Estero, Bords du Rio Salado 
La Palisa del Bracho 25 kil. N.O. DTcaiio E R. 
Wagner 1909." 

Condition. — Excellent except for loss of hind 
legs. Originally on pin, now mounted (by Ross) in 
balsam on slide. 

Description. — .Appearance: Rather large, slen- 
der, large-winged; generally pale brown to tan in 
color. Color details (dry): Cranium pale brown, 
without distinct pattern; eyes black; basal antennal 
segment concolorous with cranium, basal flagellar 
segments pale tan. distals becoming medium brown 
except for 3 white apical segments (26 segments in 
complete antenna). Thorax, legs, and abdomen con- 
colorous with cranium; wings pale tan with broad, 
weakly-defined hyaline stripes. Dimensions (on 
slide): Body length 1 1.5 mm; forewing length 9.25 
mm, breadth 2.25 mm. 

Important integumental characters. — Cranium 
short, broad; caudal margin indented on each side. 
Eyes very large, as long as head-sides behind eyes 



and broader than the cranial interspace. Mandibles 
small, oligotomoid; dentations acute, well-spaced; 
apical tooth on same plane as others, middle tooth 
of left mandible almost as large as the basal. Sub- 
mentum quadrate weakly sclerotized, no inflexed 
margins. Terminalia, except for processes and 
lobes, weakly sclerotized. Left hemitergite (10 L) 
with outer and caudal margins weak, inner-basal 
margin inflexed; process (10 LP) abruptly sclerotic, 
conspicuous, darkly pigmented, long, slender, par- 
allel-sided until twisted at apical third; arcuated at 
base, then becoming straight. Right hemitergite, 
(10 R) weakly margined except caudally, meso- 
basal side blending into cleft membrane; its process 
( 10 RP) a narrowly-tapered, sclerotic spine directed 
latero-ventrad. Medial flap (MP) obsolete, simply a 
membranous fold angled leftward into cleft. 
Epiproct (EP) large but obscure, its sclerite hori- 
zontal, weakly sclerotized; membranous surface 
partially microspiculate. Left paraproct (LPPT) 
weak at base, increasingly sclerotized caudad; 
caudo-ventral margin bearing a low, broadly-round- 
ed microspiculate nodule. Lobe of left cercus large, 
broadly expanded, surface densely echinulate. 

Additional records. — Argentina: "Ing Juarez, 




Figure 36. Ochrembia wagneri (Navas), type. Type local- 
ity: 25 km NE Icaiio, Santiago del Estero. Argentina. 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



75 



Formosa 2-7-1-1948 (R. Goldbach)." 1 male in 
Inst. Miguel Lillo, Tucuman. This specimen agrees 
closely with the above type but differs in having the 
left and caudal margins of 10 L well sclerotized 
and in the finer, more evenly and denser echinula- 
tion of the left cercus lobe. Bolivia: San Antonio de 
Parapeti, Rio Parapeti (near Camiri) 1-5-IX-1964, 
at light (B. Malkin), 3 males, CAS. These males 
closely resemble the above specimen except for the 
straight, slanted basal side of the left cercus lobe 
with its more coarse echinulation. 

Mesoamerican and Mexican 
Scelembiinae 

Except for human-introduced Pararhagadochir 
trinitatis (Ross, 1992:124), the Embiidae of these 
regions have peculiar characters found in no South 
American genus. These include a basal, or no lob- 
ing, or acute, non-echinulate distal lobing of the 
left cercus. One is tempted to speculate that dis- 
tinctions evolved during the long period in which 
the North and South American continents were 
separated. 

Because the North American genera were ex- 
tensively treated in my Mexican paper (Ross, 
1984), the following treatments are brief My 1984 
figures of the male terminalia are republished for 
the reader s convenience. 

Genus Neorhagadochir Ross 

Neorhagadochir Ross, 1944:418; 1984:4.— Szumik, 
1966:55; 1998:141 (in a cladogram). 

Type species. — Neorhagadochir inflata Ross, 
1944, by original designation. 

Distribution. — Southern Mexico to Costa Rica. 

Discussion. — This important genus is divided 
into subgenera: Neorhagadochir, s. str., for N. (N.) 
inflata Ross and Drepanembia Ross for a complex 
of species related to N. (D.) salvini (McLachlan). 
The two subgenera are readily distinguished by the 
length and shape of the left tergal process and 
many other characters. 

Neorhagadochir {N.) inflata Ross 

(Figure 37) 

Neorhagadochir inflata Ko5s, 1944:419; 1984:5. 

Holotype. — Male, on slide, USNM. Data: Gua- 



temala: Cayuga, V-15 (Wm. Schaus). This locality 
is in the lower Montagua Valley. 

Discussion. — Until very recently this easily rec- 
ognized species was known only from its holotype, 
probably collected at light. Dr. Peter W. Kovarik 
collected five males at light in Belize, Orange 
Walk, Distr, Rio Bravo during April 1995 and per- 
mitted me to retain them in CAS. 

Subgenus Drepanembia Ross 

Neorhagadochir (Drepanembia) Ross, 1984:5. 

Type species. — Embia salvini McLachlan, 
1877, by original designation. 

Distribution. — Southern Mexico to Costa Rica. 

Neorhagadochir (D.) salvini 

(McLachlan) 

(Figure 38) 

Embia salvini McLachlan, 1877:380. — Enderlein, 
1912:51. 

Embia (Olyntha) salvini McLachlan, Hagen, 1885:198. 

Olyntha salvini (McLachlan), Krauss, 191 1:31. 

Embolyntha salvini (McLachlan), Davis, 1940b:349. 

Neorhagadochir salvini (McLachlan), Ross, 1944:419; 
1984:6. 

Haploembia neosolieri Mariilo and Marquez, 1983:51, 
new synonym. 

Holotype. — Male, BMNH. Data: Guatemala: 
"Chinuatta," 4100 ft (Salvin). The type locality is a 
suburb of Guatemala City. 

Discussion. — I have cultured extensive series of 
Drepanembia from various localities in southern 
Mexico, and, during a trip by road to the subgenus" 
southernmost known occurrence in Costa Rica. In- 
cluded is a series from A^. (D.) salvini 's type local- 
ity in Guatemala. These series indicate that the sub- 
genus comprises a complex of species or sub- 
species, without striking terminalia distinctions but 
which have diff"erences in size, degree of apterism 
and coloration. For example, two unusual lots from 
seasonally arid Nicaraguan localities are subter- 
ranean and have pale amber coloration in both 
sexes. Intensive study based on these and other 
large, cultured series will be needed to properly 
treat the systematics of this subgenus. 

The description of well-figured A'. (D.) salvini 
(by Ross, 1984, and by Davis, 1940b), as a new 



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TERMINALIA 

VENTRAL 

Figure 37. Neorhagadochir {Neorhagadochir) inflata Ross, holotype (after Ross, 1984). Type locality: Cayuga, Mon- 
tagua Valley, Guatemala. 



species of the Old World genus Haploemhia by 
Marino and Marquez (1983) is most surprising. 

Genus Brachypterembia Ross 

Brachypteremhia Ross, 1984:7.— Szumik, 1998:141 (in 
a cladogram). 

Type species. — Brachypterembia moreliensis 
Ross, by original designation. 

Distribution. — Mexico: Pine-oak zone of 
mountains SE of Morelia, Michoacan, 8000 ft. 
elev. 

Discussion. — This monotypic genus is readily 
recognized by its short wings (it is expected that 
normal winged males may occur in other locali- 
ties), its long, sickle-shaped left tergal process and 
the conical basal segment of the left cercus which 
lacks a definite lobe and echinulations. 



Brachypterembia moreliensis Ross 

(Frontispiece and Figure 39) 

Brachypterembia moreliensis Ross, 1984:7. 

Holotype. — Male, on slide, CAS. Data; Mexi- 
co: Pare Nac. Jose Maria Morelos, 14 mi. SE More- 
lia, 8000 ft. elev. (E. S. Ross). 

Discussion. — This distinct species is easily rec- 
ognized by its generic characters and the figure of 
its terminalia. ft occurs commonly under pine and 
oak bark flakes and loose bark of fence posts. 

Genus Conicercembia Ross 

Conicercembia Ross, 1984:10.— Szumik, 1998:141 (in a 
cladogram). 

Conicerembia (sic), Szumik, 1996:55. 

Clothoda Enderlein, Marifio and Marquez, 1987:64 



ROSS: EMBIA, BIOSYSTEMATICSOFTHE ORX)ER EMBIIDINA, PART 3 



77 




Figure 38. Neorhagadochir (Drepanembia) salvini (McLachlan), topotype (after Ross, 1984). Type locality: Chinuatta 
(Guatemala City), Guatemala. 



Type species. — Conicercembia tepicensis Ross 
by original designation. 

Distribution. — Mexico: Sierra Madre Occiden- 
tal. 

Discussion. — This is the only Mexican embiid 
genus with a distal inner lobe on the left cercus. 
However, it may not be a homolog of a lobe in this 
position elsewhere in Embiidae because the lobe 
lacks echinulations. In fact, there are no echinula- 
tions at all on the basal cercus segment, even on the 
slight swellings at its inner base. 

The component species are very distinct; the 
type species having a sickle-shaped left tergal 
process and C. septentrionalis (Marifio and 
Marquez) having this process straight, and other 
distinctions. 



Conicercembia tepicensis Ross 

(Figure 40) 
Conicercembia tepicensis Ross, 1984:1 1. 

Holotype. — Male, on slide, CAS. Data. — Mex- 
ico: El Ocotillo, 3 mi NW of Chapahlla. Hyw. 
No. 15, 4000 ft. elev. (E. S. Ross). 

Discussion. — This species is readily identified 
by reference to the republished figure of the temii- 
nalia. The habitat is oak woodland. Most colonies 
were found under loose bark of fi"eshly installed 
fence posts. 

Conicercembia septentrionalis 

(Marifio-Marquez) 

new combination 

(Figures 4 lA-E ) 

Clothoda septentrionalis Mariiio and Marquez, 1988:64. 



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TERMINALIA 

VENTRAL 



Figure 39. Brachypterembia moreliensis Ross, holotype (after Ross, 1984). Type locality: 14 mi SE Morelia, Mexico. 



Holotype. — Male, IBUNAM. Data. — Mexico: 
Michoacan: km 37, camino Coalcoman - Las 
Nieves, 2050 m, 6 May 1983 (H. Brailovsky, E. 
Barrera and E. Marino). 

Unfortunately this species was described from a 
single male specimen with wing vein MP aberrant- 
ly forked instead of unforked. This seemingly ple- 
siomorphic character probably caused the authors 
to assign their new species to the order "s most ple- 
siomorphic genus, Clothoda Enderlein. Acmally, 
MP normally is simple in Clothoda, the multi- 
branched cubitus being its most plesiomorphic ve- 
national character. In my many specimens of what 
may be C. septentrionalis. the forking of MP is 
variable, in some it is forked in the forewings of a 
specimen but not in its hind wings. It may even be 
forked or unforked in the left or right wings of a 
single specimen. 



I have decided that Clothoda septentrionalis 
should be placed in Conicevcembia Ross. It is very 
distinct, however, from C tepicensis, the type 
species of this genus, as evidenced, for example, by 
the almost straight left tergal process (10 LP) and 
its more elongate, narrowly-acute right tergal 
process (10 RP). Additionally, a broad well-sclero- 
tized epiproct sclerite (EP) is present in C. tepicen- 
sis. whereas in C. septentrionalis this sclerite is al- 
most obsolete, being represented only by a minute 
spine beneath 10 LP. 

Series of both sexes were reared from females 
and brood collected in two localities in pine-oak 
zones of Mexico's Sierra Madre Occidental. The 
largest series is fi"om Puerto de Mazos, 1035 m 
elev. (crest of pass), about 8 miles SW of Autlan 
(Hyw. 80), Jahsco (Figs. 41A, B, D). A smaller se- 
ries was reared from stock collected about 5 miles 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



79 




Figure 40. Conicercembia tepicensis Ross, holotype (after Ross, 1984). Type locality: 3 mi NW Chapalilla, Tepic, 
Mexico. 



S. of Tecalitlan (Hwy. 110), 1240 m. elev., Colima 
(Figs. 41C, E). Adults in both cultures matured dur- 
ing May. Both occur on oak and pine bark, espe- 
cially beneath loose bark on fence posts. Each 
colony consisted of a female and her brood of early 
instar nymphs. 

It is possible that these two series represent dis- 
tinct species, as well as being distinct from C. 
septentrionalis. As figured, there are slight distinc- 
tions in the shape of tergal processes and the left 
cercus lobe. The submentum of Puerto de Mazos 
males is broader than long (that of Tecalitlan males 
quadrate), the scape and basal antennal segments 
and mandibles are amber in the former, dark brown 
in the latter. More collecting in the Sierra Madre 
Occidental may reveal existence of a complex of 
closely related species and races. 1 have taken the 
conservative approach and regard specimens from 



all three localities as representation of one variable 
species. 

INCERTAE SEDIS 

Embia (Olyntha) inuUeri Hagen 

Embia {Olvntha) miilleii Hagen, 1885:206 — Krauss, 
1911:32.— Enderlein, 1912:52.— Navas, 1918:102.— 
Davis, 1940b:352.— Ross, 1944:497 (as unrecogniz- 
able). 

Type. — Mature female, dry and crushed, MCZ. 

Type labels. — "Type 156" (red label), "Embiden 
Larve," "Itajahy Brazil 1879 Fr. Mueller." "Itajahy" 
undoubtedly is the costal town of Itajai, state of 
Santa Catarina. 

Description (based on my examination). — En- 
tire (dry) specimen black except for a narrow, 
whitish band between each thoracic somite. Crani- 



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SUBMENTUM 



Figure 41. Conicercemhia septentrionalis CMarino and Marquez). Type locality: km 37. camino Coalcoma-Las Nieves, 
Michoacan, Mexico. Figures are of specimens from northern localities: Figs. 41A, B, D based on specimen from Puer- 
to de Mazos. 8 mi SW Autlan. Jalisco; Figs.41C and E based on specimen from 5 mi. S. Tecalitlan. Colima. 



um broad, circular, rugulose in clypeal region, en- 
tire surface microrugose. dull, \estiture cream 
white. Eyes, antennae (only nine segments present), 
and palpi concolorous with cranium. Preclypeal 
membrane paler than cranium. Body sclerites 
rather smooth, glossy black; meso- and meta-scuta 
finely rugose medially; body pubescence pale yel- 
low. All legs unicolorous. dark brown, concolorous 
with body; hind basitarsi with chestnut brown plan- 
tar setae, second tarsal papilla not clear in the dry 
specimen but probably present; terminal hind tarsal 
segment black but abruptly golden in apical fifth. 
cla\\'s gray white except for dark brown distal third. 
Genital sternites nearly concolorous with other ab- 
dominal stemites. Cerci uniformly black. Body 
length: 15 mm. 

Discussion. — During early 1999 I attempted. 



without success, to collect and culture topotypic 
specimens of £. (O.)muUeri. However, because the 
type locality has greatly changed during more than 
one hundred years since the type was collected it 
may no longer be collected. It is also possible that 
the name "Itajai" may represent a region, not the 
town. 

It is likely that the species, or close relatives, 
may have wide distribution in SE Brazil for I col- 
lected females almost identical to the type oiE. (O) 
miilleri west of Curitiba but to date no males have 
mamred in the culture. Unfortunately, the culture is 
diseased for males upon becoming penultimate fail 
to complete de\elopment. Through the transparent 
derm of one dead male nymph, adult terminalia 
structures are visible, yet distorted. These struc- 
tures show that the species is generically distinct. 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



81 



The locality is on PR376 near Sao Luis Puruna, 
Balsa Nova, about 25 km W of Curitiba, 1 100 m. 
Colonies were rare in shale crevices in a high cliff 
opposite a fruit stand. Because of the much higher 
altitude, it is possible that the species isn't E. (O) 
miilleri. but almost certainly it is congeneric. 

Similar females (CAS) were collected near 
Nova Teutonia, Santa Catarina, by Fritz Plaumann. 

Pachylembiinae Ross 

new subfamily 

Type genus. — Pachylembia Ross, by present 
designation. 

Distribution. — Mexico: Sierra Madre Occiden- 
tal. 

Diagnosis. — Males apterous, nymphaform; cra- 
nium oval, eyes small, mandibles coarsely dentate 
on a single plane; submentum sclerotic, sides and 
anterior margin inflexed. Hind basitarsus stout, 
with two papillae, setae dense. Basal margins of ab- 
dominal terga 7 to 9 with increasingly large lateral 
apodemes. Tergite 10 transverse, broadly cleft; 
basal cleft margin narrowly sclerotized. Sclero- 
tized, margins of left hemitergite (10 L) inflexed, 
especially the posterior which projects caudad; 
very densely clothed with large setae which gradu- 
ally decrease in length caudad while becoming in- 
creasingly erect and stiff. Left process (10 LP) very 
small, needle-like; at times entirely concealed be- 
neath projected meso-caudad margin of 10 L. Right 
hemitergite (10 R) very broad, caudal two-thirds 
densely clothed with caudally-slanted setae, with a 
large portion of its surface depresssed, bearing very 
fine setae, or none. Right process (10 RP) usually 
spatulate and distally rounded; talon and its sub- 
tended nodule absent. Medial flap (MF) a diagonal, 
sclerotic "rod"; adjacent cleft membrane lacking a 
microspiculate depression. Epiproct sclerite (EP) 
broad, conspicuous. Hypandrium (H) very large, 
evenly sclerotized, lacking a process. Left paraproct 
(LPPT) membranous except for a narrow, medial 
sclerite. Basal segment of left cercus (LCi) largely 
membranous, lacking an innner lobe; instead, bear- 
ing either a dorso-basal, small, partly membranous, 
nipple-like nodule directed upward, or a membra- 
nous triangular flap in this position. 

Females. — ^Not investigated for subfamily char- 
acters. 

Discussion. — Pachylembiine males differ from 



all other embiids in their minute, needle-like left 
tergal process, the abundance of short-to-long, 
erect setae on both hemitergites (10 L and 10 Rj, 
and several other features. It is likely that these 
dense setae help to prolong copulatory union in 
place of well developed processes and lobes. 

The subfamily includes only its type genus 
which comprises five species. 

Genus Pachylembia Ross 

Pachylembia Ross, 1984:13.— Szumik, 1998:141 (in a 
cladogram). 

Type species. — Pachylembia chapalae Ross, by 
original designation. 

Distribution. — Mexico: Sierra Madre Occiden- 
tal. 

Diagnosis. — Characters of the subfamily. 

Discussion. — All species occur under stones 
and leaf litter at higher elevations in the Sierra 
Madre Occidental. Because both sexes are apter- 
ous, it is likely that several difficult-to-define racial 
and species complexes exist. Important characters 
include coloration and dermal texture. 



KEY TO SPECIES OF PACHYLEMBIA 

(Males) 

1. Right tergal process (10 RP) narrow, thumb- 
like. Basal segment of left cercus entirely 
membranous, dorso-basally bearing a broad, 
thin, leftward-folded flap taxcoensis 

— 10 RP broad, spatulate. Basal segment of left 
cercus only partially sclerotized, bearing 
dorso-basally a small, erect, nipple-like lobe 
2 

2. Thorax with only one narrow white band (on 
anterior margin of mesonotum) unicinta 

— Thorax with two white bands 3 

3. Cranium and all body sclerites, especially of 

the thorax, dull jet black, microrugose 

autlanae 

— Cranium and body dark brown, surface 
smooth, rather glossy 4 

4. Relatively large (body length averaging 13.0 
mm). Always uniformly dark brown. Hemiter- 
gites (10 L and 10 R) very densely setose, in- 
cluding entire caudal half of 10 R. Basal seg- 
ment of left cercus tapered caudad, its inner 
side straight chapalae 



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— Relatively small (body length averaging 10.0 
mm). Body pigmentation variable (about 25% 
of type population with cranium, mandibles 
and submentum orange). Basal segment of left 
cercus short, globose, inner side aracuate 
colimae 

Pachylembia chapalae Ross 

(Figure 42) 

Pachylembia chapalae Ross, 1984:16. 

Holotype. — Male, on slide, CAS. Data. — Mex- 
ico: Michoacan, 4 mi. NW of Sahuayo (E. S. Ross). 

Discussion. — This species, occurring in the vi- 
cinity of Lago Chapala, is fully treated in the origi- 
nal reference. Distinguishing features of males in- 
clude light to dark mahogany brown coloration, 
rather glossy surface texture; acrotergite 1 pale yel- 
low or chestnut; acrotergite 2 dark mahogany 
brown; head never golden; submentum very dark, 
poststerna blackish brown; antenna 28 segmented; 
10 L setae very dense, erect, much shorter and 
thicker on caudal margin which is very sclerotic 
and projected; 10 R longitudinally depressed in 
inner caudal area, this surface with very fine setae, 
or none. 

Pachylembia aiitlanae Ross 
new species 

Holotype. — Male, on slide, CAS. Data. — Mex- 
ico: NE of Autlan (on hyw. 80) at junction of access 
road to Microndas San Francisco. Jalisco, 1025 m; 
matured in culture 5-1-90 (E. S. Ross). 

Description. — Appearance: Rather large, apter- 
ous; entirely dull black except for two very narrow, 
white thoracic bands. Color details (in alcohol): 
Cranium very dull black, lacking pattern; clouded 
golden mahogany brown ventrally. Basal antennal 
segments dark mahogany brov\n. other segments to 
apex (segment 28) lighter mahogany brown. 
Labrum and mandibles blackish brown, other 
mouthparts mahogany brown except submentum 
which is blackish brown. All body sclerites, espe- 
cially dorsally, lusterless dull black except acroter- 
gite 1 which is brown, margined with golden 
brown; acrotergite 2 dull black. All body mem- 
branes very dark lavender except for a very narrow 
white band between each thoracic somite. All legs 
varied shades of dark, glossy, mahogany brown. 
Abdomen dull blackish brown except for slightly 




TERMINAL! A 

VENTRAL 



Figure 42. Pachylembia chapalae Ross, holotype (after 
Ross, 1984). Type locality: near Sahuayo, Michoacan, 
Mexico. 



paler membranous areas, surface with a faint metal- 
lic-blue luster; cerci dark brown except for tan non- 
sclerotic surfaces, including tips of distal segments. 
Body length 14 mm. 

Anatomical distinctions. — Cranium large, glo- 
bose; incisor teeth especially broad, wide-spaced. 
Left tergal process (10 LP) very long and slender in 
contrast to congeners, extending beyond caudal 
margin of 10 L, or equal to it. 

Allotype. — Female, in alcohol (CAS). Data. — 
Reared in holotype's culture. 

Description. — Sclerotized surfaces glossy dark 
mahogany brown except for broad, cream white 
thoracic bands, membranes otherwise purple. An- 
tennae golden tan (25 segments). Labrum dark 
amber, mandibles piceous; other mouthparts, in- 
cluding venter of cranium, golden brown. Acroter- 



ROSS: EMBIA, BIOSYSTEMATICS OF THE ORDER EMBIIDINA, PART 3 



83 



gites dark mahogany brown; thoracic sternites 
golden brown; intensity of thoracic bands increased 
by very white fat on anterior edge of acrotergite 1 
and anterior margin of mesonotum. Abdominal 
terga concolorous with thoracic scuta; sternites 
translucent amber except for brown paragenitals 
and paraprocts; cerci dark brown with pale tips. 
Body length; 16.0 mm. 

Paratypes and parallotypes. From holotype's 
culture. Deposited in CAS, USNM, and UNAM. 

Additional record. — A series from 6 mi. SW 
Union de Tula, Jalisco, 1410 m (a short distance 
SW of Autlan on Hyw. 80). 

Pachylembia colimae Ross 
new species 

(Figure 43) 

Holotype. — Male, on slide, CAS. Data. — Mex- 



ico: 15 km NE Colima, Colima, 1230 m, matured in 
culture 1-90 (E. S. Ross). 

Description. — Appearance: Medium sized, 
apterous; almost entirely dark brown except for two 
conspicuous white thoracic bands. Color details (in 
alcohol): Cranium largely piceous brown dorsally 
becoming dark golden brown between antennal 
bases and eyes, and ventrally; vertex pattern indis- 
tinct. Basal two antennal segments medium brown, 
others tan to apex (22 segments). Mandibles dark 
amber, piceous apically and basally; palpi medium 
brown; submentum dark amber, margins largely 
piceous. Thoracic sclerites varied shades of glossy, 
blackish brown, except acrotergite 1 which is very 
pale, translucent yellow, and prothoracic postemum 
which is yellow amber; all thoracic membranes 
dark lavender except when white between each tho- 
racic somite (forming two conspicuous bands); legs 
varied shades of mahogany brown. Abdominal 




TERMINALIA 

VENTRAL 
Figure 43. Pachylembia colimae Ross, holotype. Type locality: near Colima, Mexico. 



84 



OCCASIONAL PAPERS OF THE CALIFORNIA ACADEMY OF SCIENCES 



terga mahogany brown except terminalia which are 
piceous; all membranes purple; cerci weakly scle- 
rotized, light brown; extensive membranous areas, 
including tips of distal segments, pale tan. Body 
length 10.0 mm. 

Allotype. — Female, in alcohol (CAS). Data: 
Reared in holotype's culture. 

Description. — Coloration similar to that of 
males but overall paler. Cranium entirely orange 
with faint vertex pattern. Basal antennal segments 
lemon yellow, others tan to apex (22-segments). 
Body length 12.0 mm. 

Paratypes and parallotypes. — Numerous adult 
males and females reared in holotype's culture, de- 
posited in CAS, USNM, UNAM and BMNH. 

Distinguishing features. — Males are quite col- 
orful. Their body is dark, glossy chocolate brown; 
acrotergite 1 is colorless, transparent. About 25% 
of males have the cranium brilliant orange, the ma- 
jority have the head dark brown. Setae on the 
hemitergites are fewer in number than in P. cha- 
palae, less dense (sockets more widely spaced); 10 
R lacks a longitudinal inner caudal depression; 10 
LP is exceptionally minute, short, almost obsolete 
(not visible from above); 10 RP blends caudad from 
brown to amber; LCi is stubby, globose, inner mar- 
gin arcuate, dorso-basal process reduced, at times 
obsolete, (if present triangular in profile). 

Biology. — The type locality is a large rocky, al- 
most level pasture without trees. Undoubtedly it 
was once forested but perhaps cleared very soon 
after the nearby town of Colima was established. 
Colonies were very rare under the numerous vol- 
canic stones. 

Pachylembia unicincta Ross 

Pachylembia unicincta Ross, 1984:18. 

Holotype. — Male, on slide, CAS. Data. — Mex- 
ico: Jalisco. 5 km E of Mazamitla, 5800 ft. elev., 
matured in culture 1-79 (E. S. Ross). 

Discussion. — Both sexes are readily recognized 
by the presence of only one pale thoracic band in- 
stead of t\vo. Colonies were rare in oak leaf litter on 
an open slope used by picnickers in a pine-oak for- 
est adjacent to weekend cottages. 

Pachylembia taxcoensis Ross 

(Figure 44) 
Pachylembia taxcoensis Ross, 1984:14. 




Figure 44. Pachylembia taxcoensis Ross, holotype (after 
Ross, 1984). Type locality: Taxco, Gro., Mexico. 



Holotype. — Male, on slide, CAS. Data. — Mexi- 
co: Guerrero, 5 km E of Taxco, matured in culture 
20-X-76 (E. S. Ross) 

Discussion. — ^Nothing to add to the original de- 
scription. The species is easily distinguished by its 
small, narrow process of the right hemitergite (10 
RP). In congeners this process is broad, spatulate. 



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